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WORLD'S     FAMOUS     BOOKS 


The  Descent  of  Man 

And    Selection    in    Relation    to    Sex 


By 

Charles  Darwin 

Revised  Edition 


Illustrated 


New  York  and  London 
MERRILL    AND    BAKER 

Publishers 


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PREFACE  TO  THE  SECOND  EDITION. 


During  the  successive  reprints  of  tlie  first  edition  of  this  work, 
published  in  1871,  I  was  able  to  introduce  several  important  cor- 
rections; and  now  that  more  time  has  elapsed,  I  have  endeavored 
to  profit  by  the  fiery  ordeal  through  which  the  book  has  passed, 
and  have  taken  advantage  of  all  the  criticisms  which  seem  to  me 
sound.  I  am  also  greatly  indebted  to  a  large  number  of  corre- 
spondents for  the  communication  of  a  surprising  number  of  new 
facts  and  remarks.  These  have  been  so  numerous,  that  I  have 
been  able  to  use  only  the  more  important  ones;  and  of  these,  as 
well  as  of  the  more  important  corrections,  I  will  append  a  list. 
Some  new  illustrations  have  been  introduced,  and  four  of  the  old 
drawings  have  been  replaced  by  better  ones,  done  from  life  by 
Mr.  T.  W.  Wood.  I  must  especially  call  attention  to  some  observa- 
tions which  I  owe  to  the  kindness  of  Prof.  Huxley  (given  as  a  sup- 
plement at  the  end  of  Part  I.),  on  the  nature  of  the  differences  be- 
tween the  brains  of  man  and  the  higher  apes.  I  have  been  par- 
ticularly glad  to  give  these  observations,  because  during  the  last 
few  years  several  memoirs  on  the  subject  have  appeared  on  the 
Continent,  and  their  importance  has  been,  in  some  cases,  greatly 
exaggerated  by  popular  writers. 

I  may  take  this  opportunity  of  remarking  that  my 
critics  frequently  assume  that  I  attribute  all  changes  of 
corporeal  structure  and  mental  power  exclusively  to  the 
natural  selection  of  such  variations  as  are  often  called 
spontaneous;  whereas,  even  in  the  first  edition  of  the  'Origin 
of  Species,'  I  distinctly  stated  that  great  weight  must 
be  attributed  to  the  inherited  effects  of  use  and  disuse,  with 
respect  both  to  the  body  and  mind.  I  also  attributed  some  amount 
of  modification  to  the  direct  and  prolonged  action  of  changed  con- 
ditions of  life.  Some  allowance,  too,  must  be  made  for  occasional 
reversions  of  structure;  nor  must  we  forget  what  I  have  called 
"correlated"  growth,  meaning,  thereby,  that  various  parts  of  the 
organization  are  in  some  unknown  manner  so  connected,  that 
when  one  part  varies,  so  do  others;  and  if  variations  in  the  one 


Vi  PREFACE   TO   THE    SECOND   EDITION. 

are  accumulated  by  selection,  other  parts  will  be  modified.  Again, 
it  has  been  said  by  several  critics,  that  when  I  found  that  many 
details  of  structure  in  man  could  not  be  explained  through  natural 
selection,  I  invented  sexual  selection;  I  gave,  however,  a  tolerably 
clear  sketch  of  this  principle  in  the  first  edition  of  the  'Origin  of 
Species,'  and  I  there  stated  that  it  was  applicable  to  man.  This 
subject  of  sexual  selection  has  been  treated  at  full  length  in  the 
present  work,  simply  because  an  opportunity  was  here  first  afforded 
me.  I  have  been  struck  with  the  likeness  of  many  of  the  half- 
favorable  criticisms  on  sexual  selection,  with  those  which  ap- 
peared at  first  on  natural  selection;  such  as,  that  it  would  explain 
some  few  details,  but  certainly  was  not  applicable  to  the  extent 
to  which  I  have  employed  it.  My  conviction  of  the  power  of  sexual 
selection  remains  unshaken;  but  it  is  probable,  or  almost  certain, 
that  several  of  my  conclusions  will  hereafter  be  found  erroneous; 
this  can  hardly  fail  to  be  the  case  in  the  first  treatment  of  a  sub- 
ject. When  naturalists  have  become  familiar  with  the  idea  of 
sexual  selection,  it  will,  as  I  believe,  be  much  more  largely  ac- 
cepted; and  it  has  already  been  fully  and  favorably  received  by 
several  capable  judges. 

Down,  Beckenham,  Kent, 
September,  1874. 


CONTENTS. 


Introduction Pages  1-A 


PART  I. 
THE   DESCENT  OR  ORIGIN  OP  MAN. 


■j  CHAPTER  I. 

THE  EVIDENCE  OF  THE  DESCENT  OF  MAN  FROM  SOME 

LOWER  FORM. 

Nature  of  the  evidence  bearing  on  the  origin  of  man — Homologous 
structures  in  man  and  the  lower  animals — Miscellaneous  points 
of  correspondence— Development — Rudimentary  structures,  mus- 
cles, sense-organs,  hair,  hones,  reproductive  organs,  &c. — The 
bearing  of  these  three  great  classes  of  facts  on  the  origin  of  man 


•  CHAPTER  II. 

ON   THE    MANNER   OF   DEVELOPMENT    OF   MAN    FROM 
SOME  LOWER  FORM. 

Variability  of  body  and  mind  in  man — Inheritance — Causes  of 
variability— Laws  of  variation  the  same  in  man  as  in  the  lower 
animals — Direct  action  of  the  conditions  of  life — Effects  of  the  in- 
creased use  and  disuse  of  parts— Arrested  development— Rever- 
sion— Correlated  variation — Rate  of  increase— Checks  to  increase 
— Natural  selection— Man  the  most  dominant  animal  in  the  world 
— Importance  of  his  corporeal  structure — The  causes  which  have 
led  to  his  becoming  erect — Consequent  changes  of  structure — De- 
crease in  size  of  the  canine  teeth — Increased  size  and  altered 
shape   of  the  skull — ^Nakedness — Absence   of  a  tail— Defenseless 

condition    of    man 25 

vii 


Viij  CONTENTS. 

CHAPTER  III. 

COMPARISON  OF   THE   MENTAL.  POWERS  OF  MAN  AND 
THE  LOWER  ANIMALS. 

The  difference  in  mental  power  between  the  highest  ape  and  the 
lowest  savag-e,  immense— Certain  instincts  in  common— The  emo- 
tions—Curiosity—Imitation—Attention—Memory —  Imagination- 
Reason— Progressive  improvement— Tools  and  weapons  used  by 
animals— Abstraction,  Self-consciousness— Language— Sense  of 
beauty— Belief  in  God,  spiritual  agencies,  superstitions 88 


CHAPTER  IV. 

COMPARISON  OF  THE  MENTAL  POWERS  OP  MAN  AND 
THE  LOWER  ANIMALS— Continued. 

The  moral  sense— Fundamental  proposition— The  qualities  of  so- 
cial animals— Origin  of  sociability— Struggle  between  opposed  in- 
stincts—Man a  social  animal— the  more  enduring  social  instincts 
conquer  other  less  persistent  instincts— The  social  virtues  alone 
regarded  by  savages— The  self-regarding  virtues  acquired  at  a 
later  stage  of  development— The  importance  of  the  judgment  of 
the  members  of  the  same  community  on  conduct— Transmission 
of    moral    tendencies— Summary 


CHAPTER  V. 

ON  THE  DEVELOPMENT  OF  THE  INTELLECTUAL.  AND 
MORAL  FACULTIES  DURING  PRIMEVAL  AND  CIVIL- 
IZED TIMES. 

Advancement  of  the  intellectual  powers  through  natural  selec- 
tion—Importance of  imitation— Social  and  moral  faculties— Their 
development  within  the  limits  of  the  same  tribe— Natural  selec- 
tion a-s  affecting  civilized  nations— Evidence  that  civilized  na- 
tions were    once  barbarous 124 


chapteh  vl 

ON  THE  AFFINITIES  AND  GENEALOGY  OF  MAN. 

Position  of  man  in  the  animal  series— The  natural  system  genea- 
logical—Adaptive characters  of  slight  value— Various  small  points 
of  resemblance  between  man  and  the  Quadrumana— Rank  of 
man  in  the  natural  system— Birthplace  and  antiquity  of  man- 
Absence  of  fossil  connecting-links— Lower  stages  in  the  geneal- 
ogy of  man,  as  inferred,  firstly  from  his  affinities  and  secondly 
from  his  structure— Early  androgynous  condition  of  the  Verte- 
brata— Conclusion    142 


CONTENTS.  ix 

CHAPTER  VII. 

ON  THE  RACES  OF  MAN. 

The  nature  and  value  of  specific  characters— Application  to  the 
races  of  man — ^Arguments  in  favor  of,  and  opposed  to,  ranking 
the  so-called  races  of  man  as  distinct  species — Sub-species — Mon- 
og-enists  and  polygenists— Convergence  of  character — Numerous 
points  of  resemblance  in  body  and  mind  between  the  most  dis- 
tinct races  of  man — The  state  of  man  when  he  first  spread  over 
the  earth — Each  race  not  descended  from  a  single  pair — The  ex- 
tinction of  races— The  formation  of  races — The  effects  of  crossing 
— Slight  influence  of  the  direct  action  of  the  conditions  of  life- 
Slight  or  no  influence  of  natural  selection— Sexual  selection 162 


PART  II. 
SEXUAL  SELECTION. 


CHAPTER  VIII. 

PRINCIPLES  OF  SEXUAL  SELECTION. 

Secondary  sexual  characters— Sexual  selection— Manner  of  action 
—Excess  of  males— Polygamy — The  male  alone  generally  modified 
through  sexual  selection— Eagerness  of  the  male — Variability  of 
the  male — Choice  exerted  by  the  female — Sexual  compared  with 
natural  selection — Inheritance  at  corresponding  periods  of  life, 
at  corresponding  seasons  of  the  year,  and  as  limited  by  sex— Re- 
lations between  the  several  forms  of  inheritance — Causes  why  one 
sex  and  the  young  are  not  modified  through  sexual  selection — 
Supplement  on  the  proportional  numbers  of  the  two  sexes 
throughout  the  animal  kingdom— The  proportion  of  the  sexes  in 
relation  to  natural  selection 203 


CHAPTER  IX. 

SECONDARY     SEXUAL     CHARACTERS     IN     THE     LOWER 
CLASSES  OF  THE  ANIMAL  KINGDOM. 

These  characters  absent  in  the  lowest  classes— Brilliant  colors — 
Mollusca— Annelids — Crustacea,  secondary  sexual  characters 
strongly  developed;  dimorphism;  color;  characters  not  acquired 
before  maturity— Spiders,  sexual  colors  of;  stridulation  by  the 
males— Myriapoda  257 


CHAPTER  X. 

SECONDARY  SEXUAL  CHARACTERS  OF  INSECTS. 

Diversified  structures  possessed  by  the  males  for  seizing  the  fe- 
males—Differences between  the  sexes,  of  which  the  meaning  is 


3j  CONTENTS. 

Page 

not  understood— Difference  in  size  between  the  sexes— Thysanura 
— Diptera— Hemiptera— Homoptera,  musical  powers  possessed  by 
the  males  alone— Orthoptera,  musical  instruments  of  the  males, 
much  diversified  in  structure;  pugnacity;  colors— NeuVoptera  sex- 
ual differences  in  color— Hymenoptera,  pugnacity  and  colors— 
Coleoptera,  colors;  furnished  with  great  horns,  apparently  as  an 
ornament;  battles;  stridulating  organs  generally  common  to  both 
sexes  • 


272 


CHAPTER  XI. 

INSECTS,   continued.— ORDER  LEPIDOPTERA. 
(Butterflies  and  moths.) 

Courtship  of  butterflies— Battles— Ticking  noise— Colors  common  to 
both  sexes,  or  more  brilliant  in  the  males— Examples— Not  due  to 
the  direct  action  of  the  conditions  of  life— Colors  adapted  for  pro- 
tection—Colors of  moths— Display— Perceptive  powers  of  the  Lep- 
idoptera— Variability— Causes  of  the  difference  in  color  between 
the  males  and  females— Mimicry,  female  butterflies  more  bril- 
liantly colored  than  th'e  males— Bright  colors  of  caterpillars- 
Summary  and  concluding  remarks  on  the  secondary  sexual  char- 
acters of  insects— Birds  and  insects  compared 304 


CHAPTER  XII. 

SECONDARY  SEXUAL,  CHARACTERS  OF  FISHES,  AMPHIB- 
IANS, AND  REPTILES. 

Fishes:  Courtship  and  battles  of  the  males— Larger  size  of  the 
females— Males,  bright  colors  and  ornamental  appendages;  other 
strange  characters— Colors  and  appendages  acquired  by  the  males 
during  the  breeding-season  alone— Fishes  with  both  sexes  bril- 
liantly colored— Protective  colors— The  less  conspicuous  colors 
of  the  female  cannot  be  accounted  for  on  the  principle  of  protec- 
tion—Male fishes  building  nests,  and  taking  charge  of  the  ova  and 
young.  Amphibians:  Differences  in  structure  and  color  between 
the  sexes— Vocal  organs.  Reptiles:  Chelonians— Crocodiles- 
Snakes,  colors  in  some  cases  protective— Lizards,  battles  of— Or- 
namental appendages— Strange  differences  in  structure  between 
the  sexes— Colors— Sexual  differences  almost  as  great  as  with 
birds 321 


CHAPTER  XIII. 

'SECONDARY   SEXUAL,  CHARACTERS  OP  BIRDS. 

Sexual  differences— Law  of  battle— Special  weapons— Vocal  organs 
— Instrumental  music— Love-antics  and  dances— Decorations,  per- 
manent and  seasonal— Double  and  single  annual  moults — Display 
of  ornaments  by  the  males 355 


CONTENTS.  Xi 

Page 
CHAPTER  XIV. 

BIRDS.— Continued. 

Choice  exerted  by  the  female — Length  of  courtship— Unpaired  birds 
— Mental  qualities  and  taste  for  the  beautiful — Preference  or  an- 
tipathy shown  by  the  female  for  particular  males — Variability  of 
birds — Variations  sometimes  abrupt — Laws  of  variation — Forma- 
tion of  ocelli — Gradations  of  character — Case  of  Peacock,  Argus 
pheasant  and  Urosticte 399 


CHAPTER  XV. 

BIRDS.— Continued. 

Discussion  as  to  why  the  males  alone  of  some  species,  and  both 
sexes  of  others  are  brightly  colored — On  sexually-limited  inherit- 
ance, as  applied  to  various  structures  and  to  brightly-colored 
plumage — Nidification  in  relation  to  color — Loss  of  nuptual  plum- 
age during  the  winter , 438 


CHAPTER  XVI. 

BIRDS.— Concluded. 

The  immature  plumage  in  relation  to  the  character  of  the  plum- 
age in  both  sexes  when  adult — Six  classes  of  cases — Sexual  differ- 
ences between  the  males  of  closely-allied  or  representative  spe- 
cies— The  female  assuming  the  characters  of  the  male — Plumage 
of  the  young  in  relation  to  the  summer  and  winter  plumage  of 
the  adults — On  the  increase  of  beauty  in  the  birds  of  the  world — 
Protective  coloring — Conspicuously-colored  birds— Novelty  appre- 
ciated—Summary of  the  four  chapters  on  birds 458 


CHAPTER  XVII. 

SECONDARY  SEXUAL  CHARACTERS  OF  MAMMALS. 

The  law  of  battle — Special  weapons,  confined  to  the  males— Cause 
of  absence  of  weapons  in  the  female — Weapons  common  to  both 
sexes,  yet  primarily  acquired  by  the  male — Other  uses  of  such 
weapons — Their  high  importance — Greater  size  of  the  male — 
Means  of  defense — On  the  preference  shown  by  either  sex  in  the 
pairing  of  quadrupeds 496 


CHAPTER  XVIII. 

SECONDARY   SEXUAL   CHARACTERS    OP   MAMMALS— Con- 
tinued. 

Voice— Remarkable  sexual  peculiarities  in  seals— Odor— Develop- 
ment of  the  hair — Color  of  the  hair  and  skin— Anomalous  case  of 
the  female  being  more  ornamented  than  the  male — Color  and  or- 


Xii  CONTENTS. 

Page 
naments  due  to  sexual  selection— Color  acquired  for  the  sake  of 
protection— Color,  though  common  to  both  sexes,  often  due  to 
sexual  selection— On  the  disappearance  of  spots  and  stripes  in 
adult  quadrupeds — On  the  colors  and  ornaments  of  the  Quadru- 
mana— Summary  521 


PART  III. 


SEXUAL  SELECTION  IN  RELATION  TO  MAN, 
AND   CONCLUSION. 

-^  CHAPTER  XIX. 
SECONDARY  SEXUAL  CHARACTERS  OF  MAN. 

Differences  between  man  and  woman— Causes  of  such  differences, 
and  of  certain  characters  common  to  both  sexes— Law  of  battle 

-  —Differences  in  mental  powers,  and  voice— On  the  influence  of 
beauty  in  determining  the  marriages  of  mankind— Attention  paid 
by  savages  to  ornaments— Their  ideas  of  beauty  in  woman— The 
tendency   to   exaggerate    each   natural   peculiarity 551 


i  CHAPTER  XX. 

SECONDARY  SEXUAL  CHARACTERS  OF  MAN.— Continued. 

On  the  effects  of  the  continued  selection  of  women  according  to  a 
different  standard  of  beauty  in  each  race— On  the  causes  which 
interfere  with  sexual  selection  in  civilized  and  savage  nations — 
Conditions  favorable  to  sexual  selection  during  primeval  times— 
On  the  manner  of  action  of  sexual  selection  with  mankind — On 
the  women  in  savage  tribes  having  some  power  to  choose  their 
husbands— Absence  of  hair  on  the  body,  and  development  of  the 
beard— Color  of  the  skin— Summary 580 


^^  CHAPTER  XXI. 

GENERAL  SUMMARY  AND  CONCLUSION. 

Main  conclusion  that  man  is  descended  from  some  lower  form- 
Manner  of  development— Genealogy  of  man— Intellectual  and 
moral  faculties— Sexual  selection— Concluding  remarks 601 

Index 615 


THE  DESCENT  OF  MAN; 


AND 


SELECTION  IN  RELATION  TO  SEX. 


INTRODUCTION. 

The  nature  of  the  following  work  will  be  best  understood  by  a 
brief  account  of  how  it  came  to  be  written.  During  many  years 
I  collected  notes  on  the  origin  or  descent  of  man,  without  any 
intention  of  publishing  on  the  subject,  but  rather  with  the  de- 
termination not  to  publish,  as  I  thought  that  I  should  thus  only 
add  to  the  prejudices  against  my  views.  It  seemed  to  me  sufla- 
cient  to  indicate,  in  the  first  edition  of  my  'Origin  of  Species,'  that 
by  this  work  "light  would  be  thrown  on  the  origin  of  man  and 
"his  history;"  and  this  implies  that  man  must  be  included  with 
other  organic  beings  in  any  general  conclusion  respecting  his 
manner  of  appearance  on  this  earth.  Now  the  case  wears  a 
wholly  different  aspect.  When  a  naturalist  like  Carl  Vogt  ven- 
tures to  say  in  his  address  as  President  of  the  National  Institution 
of  Geneva  (1869),  "personne,  en  Europe  au  moins,  n'ose  plus  sou- 
"tenir  la  creation  independante  et  de  toutes  pieces,  des  especes," 
it  is  manifest  that  at  least  a  large  number  of  naturalists  must  ad- 
mit that  species  are  the  modified  descendants  of  other  species; 
and  this  especially  holds  good  with  the  younger  and  rising  natural- 
ists. The  greater  number  accept  the  agency  of  natural  selection; 
though  some  urge,  whether  with  justice  the  future  must  decide, 
that  I  have  greatly  overrated  its  importance.  Of  the  older  and 
honored  chiefs  in  natural  science,  many  unfortunately  are  still 
opposed  to  evolution  in  every  form. 

In  consequence  of  the  views  now  adopted  by  most  naturalists, 
and  which  will  ultimately,  as  in  every  other  case,  be  followed  by 
others  who  are  not  scientific,  I  have  been  led  to  put  together  my 
notes,  so  as  to  see  how  far  the  general  conclusions  arrived  at  in 
my  former  works  were  applicable  to  man.  This  seemed  all  the 
more  desirable,  as  I  had  never  deliberately  applied  these  views  to 
a  species  taken  singly.  When  we  confine  our  attention  to  any 
one  form,  we  are  deprived  of  the  weighty  arguments  derived  from 
2 


2  INTRODUCTION. 

tlie  nature  of  the  affinities  which  connect  together  v/hole  groups 
of  organisms — their  geographical  distribution  in  past  and  present 
times,  and  their  geological  succession.  The  homological  struc- 
ture, embryological  development,  and  rudimentary  organs  of  a 
species  remain  to  be  considered,  whether  it  be  man  or  any  other 
animal,  to  which  our  attention  may  be  directed;  but  these  great 
classes  of  facts  afford,  as  it  appears  to  me,  ample  and  conclusive 
evidence  in  favor  of  the  principle  of  gradual  evolution.  The 
strong  support  derived  from  the  other  arguments  should,  however, 
always  be  kept  before  the  mind. 

The  sole  object  of  this  work  is  to  consider,  firstly,  whether 
man,  like  every  other  species,  is  descended  from  some  pre-exist- 
ing form;  secondly,  the  manner  of  his  development;  and  thirdly, 
the  value  of  the  differences  between  the  so-called  races  of  man. 
As  I  shall  confine  mj'^self  to  these  points,  it  will  not  be  necessary 
to  describe  in  detail  the  differences  between  the  several  races — an 
enormous  subject  which  has  been  fully  discussed  in  many  valu- 
able works.  The  high  antiquity  of  man  has  recently  been  demon- 
strated by  the  labors  of  a  host  of  eminent  men,  beginning  with 
M.  Boucher  de  Perthes;  and  this  is  the  indispensable  basis  for 
understanding  his  origin.  I  shall,  therefore,  take  this  conclusion 
for  granted,  and  may  refer  my  readers  to  the  admirable  treatises 
of  Sir  Charles  Lyell,  Sir  John  Lubbock,  and  others.  Nor  shall 
I  have  occasion  to  do  more  than  to  allude  to  the  amount  of  differ- 
ence between  man  and  the  anthropomorphous  apes;  for  Prof. 
Huxley,  in  the  opinion  of  most  competent  judges,  has  conclusively 
shown  that  in  every  visible  character  man  differs  less  from  the 
higher  apes,  than  these  do  from  the  lower  members  of  the  same 
order  of  Primates. 

This  work  contains  hardly  any  original  facts  in  regard  to  man; 
but  as  the  conclusions  at  which  I  arrived,  after  drawing  up  a 
rough  draft,  appeared  to  me  interesting,  I  thought  that  they  might 
interest  others.  It  has  often  and  confidently  been  asserted,  that 
man's  origin  can  never  be  known:  but  ignorance  more  frequently 
begets  confidence  than  does  knowledge:  it  is  those  who  know  lit- 
tle, and  not  those  who  know  much,  who  so  positively  assert  that 
this  or  that  problem  will  never  be  solved  by  science.  The  con- 
clusion that  man  is  the  co-descendant  with  other  species  of  some 
ancient,  lower,  and  extinct  form,  is  not  in  any  degree  new.  La- 
marck long  ago  came  to  this  conclusion,  which  has  lately  been 
maintained  by  several  eminent  naturalists  and  philosophers;  for 
instance,  by  Wallace,  Huxley,  Lyell,  Vogt,  Lubbock,  Biichner, 
Rolle,  &c.,^  and  especially  by  Hackel.    This  last  naturalist,  besides 

1  As  the  works  of  the  first-named  authors  are  so  well  known,  I  need 
not  give  the  titles;  but  as  those  of  the  latter  are  less  well  known  in 
Eng-land,  I  will  give  them :— 'Sechs  Vorlesungen  uber  die  Darwin'sche 


INTRODUCTION.  3 

Ms  great  work,  'Generelle  Morphologie'  (1866),  has  recently  (1868, 
with  a  second  edit,  in  1870),  published  his  'Natiirliche  Schop- 
fungsgeschichte,'  in  which  he  fully  discusses  the  genealogy  of 
man.  If  this  work  had  appeared  before  my  essay  had  been  writ- 
ten, I  should  probably  never  have  completed  it.  Almost  all  the 
conclusions  at  which  I  have  arrived  I  find  confirmed  by  this  natur- 
alist, whose  knowledge  on  many  points  is  much  fuller  than  mine. 
V/herever  I  have  added  any  fact  or  view  from  Prof.  Hackel's 
writings,  I  give  his  authority  in  the  text;  other  statements  I 
leave  as  they  originally  stood  in  my  manuscript,  occasionally  giv- 
ing in  the  foot-notes  references  to  his  workfe,  as  a  confirmation  of 
the  more  doubtful  or  interesting  points. 

During  many  years  it  has  seemed  to  me  highly  probable  that 
sexual  selection  has  played  an  important  part  in  differentiating 
the  races  of  man;  but  in  ray  'Origin  of  Species'  (first  edition,  p. 
199)  I  contented  myself  by  merely  alluding  to  this  belief.  When 
I  came  to  apply  this  view  to  man,  I  found  it  indispensable  to  treat 
the  whole  subject  in  full  detail.-  Consequently  the  second  part  of 
the  present  work,  treating  of  sexual  selection,  has  extended  to  an 
inordinate  length,  compared  with  the  first  part;  but  this  could  not 
be  avoided. 

I  had  intended  adding  to  the  present  volumes  an  essay  on  the 
expression  of  the  various  emotions  by  man  and  the  lower  animals. 
My  attention  was  called  to  this  subject  many  years  ago  by  Sir 
Charles  Bell's  admirable  work.  This  illustrious  anatomist  main- 
tains that  man  is  endowed  with  certain  muscles  solely  for  the 
sake  of  expressing  his  emotions.  As  this  view  is  obviously  op- 
posed to  the  belief  that  man  is  descended  from  some  other  and 
lower  form,  it  was  necessary  for  me  to  consider  it.  I  likewise 
wished  to  ascertain  how  far  the  emotions  are  expressed  in  the 
same  manner  by  the  different  races  of  man.  But  owing  to  the 
length  of  the  present  work,  I  have  thought  it  better  to  reserve  my 
essay  for  separate  publication. 


Theorie:'  zweite  Auflage,  1868,  von  Dr.  L.  Buchner;  translated  into 
French  under  the  title  'Conferences  sur  la  Theorie  Darwinienne,'  1869, 
'Der  Mensch,  im  Lichte  der  Darwin' sche  Lehre,'  1865,  von  Dr.  F. 
Rolle.  I  will  not  attempt  to  give  references  to  all  the  authors  who  have 
taken  the  same  side  of  the  question.  Thus  G.  Canestrini  has  published 
CAnnuario  della  Soc.  d.  Nat.,'  Modena,  1867,  p.  81)  a  very  curious 
paper  on  rudimentary  characters,  as  bearing  on  the  origin  of  maji. 
Another  work  has  (1869)  been  published  by  Dr.  Francesco  Barrage, 
bearing  in  Italian  the  title  of  "Man  made  in  the  image  of  God,  was 
"also  made  in  the  image  of  the  ape." 

2  Prof.  Hackel  was  the  only  author  who,  at  the  time  when  this  work 
first  appeared,  had  discussed  the  subject  of  sexual  selection,  and  had 
seen  its  full  importance,  since  the  publication  of  the  'Origin' ;  and  this 
iie  did  in  a  very  able  manner  in  his  various  works. 


PART  I. 

THE  DESCENT  OR  ORIGIN  OF  MAN. 


CHAPTER  I. 


THE  EVIDENCE  OF  THE  DESCENT  OF  MAN  FROM 
SOME  LOWER  FORM. 

Nature  of  the  evidence  bearing  on  the  origin  of  man— Homologous 
structures  in  man  and  the  lower  animals — Miscellaneous  points  of 
correspondence — Development— Rudimentary  structures,  muscles, 
sense-organs,  hair,  bones,  reproductive  organs,  &c. — The  bearing  of 
these  three  great  classes  of  facts  on  the  origin  of  man. 

He  who  wishes  to  decide  whether  man  is  the  modified  descend- 
ant of  some  pre-existing  form,  would  probably  first  enquire 
whether  man  varies,  however  slightly,  in  bodily  structure  and  in 
mental  faculties;  and  if  so,  whether  the  variations  are  trans- 
mitted to  his  offspring  in  accordance  with  the  laws  which  prevail 
with  the  lower  animals./  Again,  are  the  variations  the  result,  as 
far  as  our  ignorance 'permits  us  to  judge,  of  the  same  general 
causes,  and  are  they  governed  by  the  same  general  laws,  as  in  the 
case  of  other  organisms;  for  instance,  by  correlation,  the  inherited 
effects  of  use  and  disuse,  &c.?  /Is  man  subject  to  similar  mal- 
conformations,  the  result  of  arrested  development,  of  reduplica- 
tion of  parts,  &;c.,  and  does  he  display  in  any  of  his  anomalies 
reversion  to  some  former  and  ancient  type  of  structure  ?  It  might 
also  naturally  be  enquired  whether  man,  like  so  many  other  ani- 
mals, has  given  rise  to  varieties  and  sub-races,  differing  but 
slightly  from  each  other,  or  to  races  differing  so  much  that  they 
must  be  classed  as  doubtful  species?  How  are  such  races  dis- 
tributed over  the  world;  and  how,  when  crossed,  do  they  react 
on  each  other  in  the  first  and  succeeding  generations?  And  so 
with  many  other  points. ' 

The  enquirer  would  next  come  to  the  important  point  whether 
man  tends  to  increase  at  so  rapid  a  rate,  as  to  lead  to  occasional 
2 


6  THE  DESCENT  OF  MAN. 

severe  struggles  for  existence;  and  consequently  to  beneficial  vari- 
ations, whether  in  body  or  mind,  being  preserved,  and  injurious 
ones  eliminated.  Do  the  races  or  species  of  men,  whichever  term 
may  be  applied,  encroach  on  and  replace  one  another,  so  that 
some  finally  become  extinct?  Yv^e  shall  see  that  all  these  ques- 
tions, as  indeed  is  obvious  in  respect  to  most  of  them,  must  be  an- 
swered in  the  afiirmative,  in  the  same  manner  as  with  the  lower 
animals.  But  the  several  considerations  just  referred  to  may 
be  conveniently  deferred  for  a  time:  and  we  will  first  see  how 
far  the  bodily  structure  of  man  shows  traces,  more  or  less  plain, 
of  his  descent  from  some  lower  form.  In  succeeding  chapters  the 
mental  powers  of  man,  in  comparison  with  those  of  the  lower 
animals,  will  be  considered. 

The  Bodily  Structure  of  Man.—U  is  notorious  that  man  is  con- 
structed on  the  same  general  type  or  model  as  other  mammals. 
All  the  bones  in  his  skeleton  can  be  compared  with  correspond- 
ing bones  in  a  monkey,  bat,  or  seal.  So  it  is  with  his  muscles, 
nerves,  blood-vessels  and  internal  viscera.  The  brain,  the  most 
important  of  all  the  organs,  follows  the  same  law,  as  shown  by 
Huxley  and  other  anatomists.  Bischoff,^  who  is  a  hostile  witness, 
admits  that  every  chief  fissure  and  fold  in  the  brain  of  man  has 
its  analogy  in  that  of  the  orang;  but  he  adds  that  at  no  period  of 
development  do  their  brains  perfectly  agree;  nor  could  perfect 
agreement  be  expected,  for  otherwise  their  mental  powers  would 
have  been  the  same.  Vulpian^  remarks:  "Les  differences  reelles 
"qui  existent  entre  I'encephale  de  I'homme  et  celui  des  singes 
"superieurs,  sont  bien  minimes.  II  ne  faut  pas  se  faire  d'illusions 
"a  cet  egard.  L'homme  est  bien  plus  pres  des  singes  anthropomor- 
"phes  par  les  caracteres  anatomiques  de  son  cerveau  que  ceux-ci 
"ne  le  sont  nonseulement  des  autres  mammiferes,  mais  meme  de 
"certains  quadrumanes,  des  guenons  et  des  macaques."  But  it 
would  be  superfluous  here  to  give  further  details  on  the  corre- 
spondence between  man  and  the  higher  mammals  in  the  structure 
of  the  brain  and  all  other  parts  of  the  body. 

It  may,  however,  be  wortlji  while  to  specify  a  few  points,  not 
directly  or  obviously  connected  with  structure,  by  which  this 
correspondence  or  relationship  is  well  shown. 

Man  is  liable  to  receive  from  the  lower  animals,  and  to  com- 
municate to  them,  certain  diseases,  as  hydrophobia,  variola,  the 

1  'Grosshirnwindungen  des  Ivlenschen,'  1868,  s.  96.  The  conclusions  of 
this  author,  as  well  as  those  of  Gratiolet  and  Aeby,  concerning  the 
brain,  will  be  discussed  by  Prof.  Huxley  in  the  Appendix  alluded  to  in 
the  Preface  to  this  edition. 

2  'Lee.  sur  la  Phys.'  1866,  p.  890,  as  quoted  by  M.  Dally,  'L'Ordre  des 
Prinaates  ©t  le  Transformisme,'  1868,  p.  29. 


HOMOLOGICAL     STRUCTURES.  7 

glanders,  syphilis,  cholera,  herpes,  &c.;^  and  this  fact  proves  the 
close  similarity*  of  their  tissues  and  blood,  both  in  minute  struc- 
ture and  composition,  far  more  plainly  than  does  their  comparison 
under  the  best  microscope,  or  by  the  aid  of  the  best  chemical 
analysis.  Monkeys  are  liable  to  many  of  the  same  non-contagious 
diseases  as  we  are;  thus  Rengger,^  who  carefully  observed  for  a 
long  time  the  Cebus  Azarae  in  its  native  land,  found  it  liable  to 
catarrh,  with  the  usual  symptoms,  and  which,  when  often  recur- 
rent, led  to  consumption.  These  monkeys  suffered  also  from 
apoplexy,  inflammation  of  the  bowels,  and  cataract  in  the  eye. 
The  younger  ones  when  shedding  their  milk-teeth  often  died  from 
fever.  Medicines  produced  the  same  effect  on  them  as  on  us.  Many 
kinds  of  monkeys  have  a  strong  taste  for  tea,  coffee,  and  spirit- 
uous liquors:  they  will  also,  as  I  have  myself  seen,  smoke  to- 
bacco with  pleasure.^  Brehm  asserts  that  the  natives  of  north- 
eastern Africa  catch  the  wild  baboons  by  exposing  vessels  with 
strong  beer,  by  which  they  are  made  drunk.  He  has  seen  some 
of  these  animals,  which  he  kept  in  confinement,  in  this  state;  and 
he  gives  a  laughable  account  of  their  behavior  and  strange 
grimaces.  On  the  following  morning  they  were  very  cross  and 
dismal;  they  held  their  aching  heads  with  both  hands,  and  wore 
a  most  pitiable  expression:  when  beer  or  wine  was  offered  them, 
they  turned  away  with  disgust,  but  relished  the  juice  of  lemons.^ 
An  American  monkey,  an  Ateles,  after  getting  drunk  on  brandy, 
would  never  touch  it  again,  and  thus  was  wiser  than  many  men. 
These  trifling  facts  prove  how  similar  the  nerves  of  taste  must 
be  in  monkeys  and  man,  and  how  similarly  their  whole  nervous 
system  is  affected. 
Man  is  infested  with  internal  parasites,  sometimes  causing  fatal 

3  Dr.  W.  Lauder  Lindsay  has  treated  this  subject  at  some  length  in 
the  'Journal  of  Mental  Science,'  July,  1871;  and  in  the  'Edinburgh 
Veterinary  Review,'  July,  1858. 

*  A  Reviewer  has  criticized  ('British  Quarterly  Review,'  Oct.  1st,  1871, 
p.  472)  what  I  have  here  said  with  much  severity  and  contempt;  but 
as  I  do  not  use  the  term  identity,  I  cannot  see  that  I  am  greatly  in 
error.  There  appears  to  me  a  strong  analogy  between  the  same 
infection  or  contagion  producing  the  same  result  or  one  closely  similar, 
in  two  distinct  animals,  and  the  testing  of  two  distinct  fluids  by  the 
same  chemical  reagent. 

5  'Naturgeschichte  der  Saugethiere  von  Paraguay,'  1830,  s.  50. 

^  The  same  tastes  are  common  to  some  animals  much  lower  in  the 
scale.  Mr.  A.  Nicols  informs  me  that  he  kept  in  Queensland,  in  Aus- 
tralia, three  individuals  of  the  Phaseolarctus  cinereus;  and  that,  with- 
out having  been  taught  in  any  way,  they  acquired  a  strong  taste  for 
rum,  and  for  smoking  tobacco. 

7  Brehm,  'Thierleben,'  B.  i.  1864,  s.  75,  86.  On  the  Ateles,  s.  105.  For 
other  analogous  statements,  see  s.  25,  107. 


S  THE  DESCENT  OF  MAN. 

effects;  and  is  plagued  by  external  parasites,  all  of  which  belong 
to  the  same  genera  or  families  as  those  infesting  other  mam- 
mals, and  in  the  case  of  scabies  to  the  same  species.^  Man  is 
subject,  like  other  mammals,  birds,  and  even  insects,"  to  that  mys- 
terious law,  which  causes  certain  normal  processes,  such  as  ges- 
tation, as  well  as  the  maturation  and  duration  of  various  diseases, 
to  follow  lunar  periods.  His  wounds  are  repaired  by  the  same 
process  of  healing;  and  the  stumps  left  after  the  amputation  of 
his  limbs,  especially  during  an  early  embryonic  period,  occasion- 
ally possess  some  power  of  regeneration,  as  in  the  lowest  ani- 
mals.^° 

The  whole  process  of  that  most  important  function,  the  repro- 
duction of  the  species,  is  strikingly  the  same  in  all  mammals, 
from  the  first  act  of  courtship  by  the  male,"  to  the  birth  and 
nurturing  of  the  young.  Monkeys  are  born  in  almost  as  helpless 
a  condition  as  our  own  infants;  and  in  certain  genera  the  young 
differ  fully  as  much  in  appearance  from  the  adults,  as  do  our 
children  from  their  full-grown  parents.^  It  has  been  urged  by 
some  writers,  as  an  important  distinction,  that  with  man  the 
young  arrive  at  maturity  at  a  much  later  age  than  with  any  other 
animal:  but  if  we  look  to  the  races  of  mankind  which  inhabit 
tropical  countries  the  difference  is  not  great,  for  the  orang  is 
believed  not  to  be  adult  till  the  age  of  from  ten  to  fifteen  years." 
Man  differs  from  woman  in  size,  bodily  strength,  hairiness,  &c., 

8  Dr.  W.  Lauder  Lindsay,  'Edinburgh  Vet.  Review,'  July  1858,  p.  13. 

8  With  respect  to  insects  see  Dr.  Laycock,  "On  a  General  Law  of 
Vital  Periodicity,"  'British  Association,'  1842.  Dr.  MaccuUoch,  'Silli- 
man's  North  American  Journal  of  Science,'  vol.  xvli.  p.  305,  has  seen  a 
dog  suffering-  from  tertian  ague.  Hereafter  I  shall  return  to  this  sub- 
ject. 

10 1  have  given  the  evidence  on  this  head  in  my  'Variation  of  Animals 
and  Plants  under  Domestication,'  vol.  ii.  p.  15,  and  more  could  be  added. 

11  "Mares  e  diversis  generibus  Quadrumanorum  sine  dubio  dignoscunt 
"feminas  humanas  a  maribus.  Primum,  credo,  odoratu,  postea  as- 
"pectu.  Mr.  Youatt,  qui  diu  in  Hortis  Zoologicis  (Bestiariis)  medicus 
"animalium  erat,  vir  in  rebus  observandis  oautus  et  sagax,  hoc  mihi 
"certissime  probavit,  et  curatores  ejusdem  loci  et  alii  e  ministris 
"confirmaverunt.  Sir  Andrew  Smith  et  Brehm  notabant  iden  in  Cyno- 
"cephalo.  Illustrissimus  Cuvier  etiam  narrat  multa  de  hac  re,  qua  ut 
"opinor,  nihil  turpius  potest  indicari  inter  omnia  hominibus  et  Quad- 
"rumanis  communia.  Narrat  enim  Cynocephalum  quendam  in  furorem 
"incidere  aspectu  feminarum  aliquarum,  sed  nequaquam  accendi  tanto 
"furore  ab  omnibus.  Semper  eligebat  juniores,  et  dignoscebat  in 
"turba,  et  advocabat  voce  gestuque." 

12  This  remark  is  made  with  respect  to  Cynocephalus  and  the  an- 
thropomorphous apes  by  Geoffroy  Saint-Hilaire  and  F.  Cuvier,  'Hist. 
Nat.  des  Mammiferes,'  tom.  i.  1824.  , 

13  Huxley,  'Man's  Place  in  Nature,'  1863,  p.  34. 


HOMOLOGICAL    STRUCTURES.  S 

as  well  as  in  mind,  in  the  same  manner  as  do  the  two  sexes  of 
many  mammals.  So  that  the  correspondence  in  general  structure, 
in  the  minute  structure  of  the  tissues,  in  chemical  composition 
and  in  constitution,  between  man  and  the  higher  animals,  es- 
pecially the  anthropomorphous  apes,  is  extremely  close. 

Emhryonic  Development. — Man  is  developed  from  an  ovule, 
about  the  125th  of  an  inch  in  diameter,  which  differs  in  no  respect 
from  the  ovules  of  other  animals.  The  embryo  itself  at  a  very 
early  period  can  hardly  be  distinguished  from  that  of  other  mem- 
bers of  the  vertebrate  kingdom.  At  this  period  the  arteries  run 
in  arch-like  branches,  as  if  to  carry  the  blood  to  branchiae  which 
are  not  present  in  the  higher  vertebrata,  though  the  slits  on  the 
sides  of  the  neck  still  remain  (f,  g,  fig.  1),  marking  their  former 
position.  At  a  somewhat  later  period,  when  the  extremities  are  de- 
veloped, "the  feet  of  lizards  and  mammals,"  as  the  illustrious  Von 
Baer  remarks,  "the  wings  and  feet  of  birds,  no  less  than  the  hands 
"and  feet  of  man,  all  arise  from  the  same  fundamental  form."  It 
is,  says  Prof.  Huxley,^*  "quite  in  the  later  stages  of  development 
"that  the  young  human  being  presents  marked  differences  from 
"the  young  ape,  while  the  latter  departs  as  much  from  the  dog  in 
"its  developments,  as  the  man  does.  Startling  as  this  last  asser- 
"tion  may  appear  to  be,  it  is  demonstrably  true." 

As  some  of  my  readers  may  never  have  seen  a  drawing  of  an 
embryo,  I  have  given  one  of  man  and  another  of  a  dog,  at  about 
the  same  early  stage  of  development,  carefully  copied  from  two 
works  of  undoubted  accuracy.^" 

After  the  foregoing  statements  made  by  such  high  authorities, 
it  would  be  superfluous  on  my  part  to  give  a  number  of  borrowed 
details,  showing  that  the  embryo  of  man  closely  resembles  that 
of  other  mammals.  It  may,  however,  be  added,  that  the  human 
embryo  likewise  resembles  certain  low  forms  when  adult  in  vari- 
ous points  of  structure.  For  instance,  the  heart  at  first  exists  as  a 
simple  pulsating  vessel;  the  excreta  are  voided  through  a  cloacal 
passage;  and  the  os  coccyx  projects  like  a  true  tail,  "extending 

1*  'Man's  Place  in  Nature,'  1863,  p.  67. 

15  The  human  embryo  (upper  flg-.)  is  from  Ecker,  'Icones  Phys.,'  1851- 
1859,  tab.  XXX.  fig.  2.  This  embryo  was  ten  lines  in  length,  so  that  the 
drawing  is  much  magnified.  The  embryo  of  the  dog  is  from  Bischoff, 
'Entwicklungsgeschichte  des  Hunde-Eies,'  1845,  tab.  xi.  fig.  42  b.  This 
drawing  is  five  times  magnified,  the  embryo  being  twenty-five  days  old. 
The  internal  viscera  have  been  omitted,  and  the  uterine  appendages 
in  both  drawings  removed.  I  was  directed  to  these  figures  by  Prof. 
Huxley,  from  whose  work,  'Man's  Place  in  Nature,'  the  idea  of  giving 
them  was  taken.  Hackel  has  also  given  analogous  drawings  in  his 
'Schopfungsgesohichte.* 


JO 


THE  DESCENT  OP  MAN. 


Fig.  !•    Upper  figure  human  embryo,  from  Ecker.    Lower  figure  that 
of  a  dog,  from  Bischoff. 

a„  Fore-brain,  cerebral  hemispheres,  &c. 

b.  Mid-brain,  corpora  quadrigemina. 

c.  Hind-brain,  cerebellum,  medulla  oblongata. 

d.  Eye. 

e.  Ear. 

f.  First  visceral  arch. 

g.  Second  visceral  arch. 

H.  "Vertebral  columns  and  muscles  process  of  development, 
i.  Anterior  extremity. 
K.  Posterior  extremity. 
L.  Tail  or  os  coccyx. 


RUDIMENTS.  11 

"considerably  beyond  the  rudimentary  legs.""  In  the  embryos  of 
all  air-breathing  vertebrates,  certain  glands,  called  the  corpora 
Wolffiana,  correspond  with,  and  act  like  the  kidneys  of  mature 
fishes."  Even  at  a  later  embryonic  period,  some  striking  resem- 
blances between  man  and  the  lower  animals  m.ay  be  observed. 
Bischoff  says  that  the  convolutions  of  the  brain  in  a  human  foetus 
at  the  end  of  the  seventh  month  reach  about  the  same  stage  of 
development  as  in  a  baboon  when  adult."  The  great  toe,  as 
Prof.  Owen  remarks,^''  "which  forms  the  fulcrum  when  standing 
"or  walking,  is  perhaps  the  most  characteristic  peculiarity  in  the 
"human  structure;"  but  in  an  embryo,  about  an  inch  in  length, 
Prof.  Wyman-^  found  "that  the  great  toe  was  shorter  than  the 
"others;  and,  instead  of  being  parallel  to  them,  projected  at  an 
"angle  from  the  side  of  the  foot,  thus  corresponding  with  the 
"permanent  condition  of  this  part  in  the  quadrumana."  I  will 
conclude  with  a  quotation  from  Huxley,-^  who  after  asking,  does 
man  originate  in  a  different  way  from  a  dog,  bird,  frog  or  fish? 
says,  "the  reply  is  not  doubtful  for  a  moment;  without  question, 
"the  mode  of  origin,  and  the  early  stages  of  the  development  of 
"man,  are  identical  with  those  of  the  animals  immediately  below 
"him  in  the  scale:  without  a  doubt  in  these  respects,  he  is  far 
"nearer  to  apes  than  the  apes  are  to  the  dog." 

Rudiments.— This  subject,  though  not  intrinsically  more  im- 
portant than  the  two  last,  will  for  several  reasons  be  treated  here 
more  fully.™  Not  one  of  the  higher  animals  can  be  named  which 
does  not  bear  some  part  in  a  rudimentary  condition;  and  man 
forms  no  exception  to  the  rule.  Rudimentary  organs  must  be  dis- 
tinguished from  those  that  are  nascent;  though  in  some  cases  the 
distinction  is  not  easy.  The  former  are  either  absolutely  useless, 
such  as  the  mammse  of  male  quadrupeds,  or  the  incisor  teeth  of 
ruminants  which  never  cut  through  the  gums;  or  they  are  of  such 
slight  service  to  their  present  possessors,  that  we  can  hardly  sup- 
pose that  they  were  developed  under  the  conditions  which  now  ex- 

i«  Prof.   Wyman   in   'Proc.  of  American  Acad,   of  Sciences,'  vol.  iv. 
1860,  p.  17. 

1'^  Owen,  'Anatomy  of  Vertebrates,'  vol,  i.  p.  533. 

18  'Die  Grosshirnwindungen  des  Menschen,'  1868,  s.  95. 

18  'Anatomy  of  Vertebrates,'  vol.  ii.  p.  553. 

20  'Proc.  Soc.  Nat.  Hist.'  Boston,  1863,  vol.  ix.  p.  185. 

'^  'Man's  Place  in  Nature,'  p.  65. 

22  1  had  written  a  rough  copy  of  this  chapter  before  reading  a  valu*.' 
able  paper,  "Caratteri  rudimentali  in  ordine  all'  origine  del  uomo'* 
('Annuario  della  Soc,  d,  Nat.,'  Modena,  1867,  p.  81),  by  G.  Canestrini, 
to  which  paper  I  am  considerably  indebted.  Hackel  has  given  ad- 
mirable discussions  on  this  whole  subject,  under  the  title  of  Dysteleo- 
logy,  in  his   'Generelle  Morphologie'   and  'Schopfungsgeschichte.' 


12  THE  DESCENT  OF  MAN. 

ist.  Organs  in  this  latter  state  are  not  strictly  rudimentary,  but 
they  are  tending  in  this  direction.  Nascent  organs,  on  the  other 
hand,  though  not  fully  developed,  are  of  high  service  to  their  pos- 
sessors, and  are  capable  of  further  development.  Rudimentary 
organs  are  eminently  variable;  and  this  is  partly  intelligible,  as 
they  are  useless,  or  nearly  useless,  and  consequently  are  no  longer 
subjected  to  natural  selection.  They  often  become  wholly  sup- 
pressed. When  this  occurs,  they  are  nevertheless  liable  to  oc- 
casional reappearance  through  reversion — a  circumstance  well 
worthy  of  attention. 

The  chief  agents  in  causing  organs  to  become  rudimentary 
seem  to  have  been  disuse  at  that  period  of  life  when  the  organ  is 
chiefly  used  (and  this  is  generally  during  maturity),  and  also 
inheritance  at  a  corresponding  period  of  life.  The  term  "disuse" 
does  not  relate  merely  to  the  lessened  action  of  muscles,  but  in- 
cludes a  diminished  flow  of  blood  to  a  part  or  organ,  from  being 
subjected  to  fewer  alternations  of  pressure,  or  from  becoming  in 
any  way  less  habitually  active.  Rudiments,  however,  may  occur 
in  one  sex  of  those  parts  which  are  normally  present  in  the 
other  sex;  and  such  rudiments,  as  we  shall  hereafter  see,  have 
often  originated  in  a  way  distinct  from  those  here  referred  to.  In 
some  cases,  organs  have  been  reduced  by  means  of  natural  selec- 
tion, from  having  become  injurious  to  the  species  under  changed 
habits  of  life.  The  process  of  reduction  is  probably  often  aided 
through  the  two  principles  of  compensation  and  economy  of 
growth;  but  the  later  stages  of  reduction,  after  disuse  has  done 
all  that  can  fairly  be  attributed  to  it,  and  when  the  saving  to  be 
effected  by  the  economy  of  growth  would  be  very  small,^^  are  diffi- 
cult to  understand.  The  final  and  complete  suppression  of  a  part, 
already  useless  and  much  reduced  in  size,  in  which  case  neither 
compensation  nor  economy  can  come  into  play,  is  perhaps  intelli- 
gible by  the  aid  of  the  hypothesis  of  pangenesis.  But  as  the 
whole  subject  of  rudimentary  organs  has  been  discussed  and 
illustrated  in  my  former  works,^*  I  need  here  say  no  more  on  this 
head. 

Rudiments  of  various  muscles  have  been  observed  in  many 
parts  of  the  human  body;=^  and  not  a  few  muscles,  which  are  regu- 

23  Some  g-ood  criticisms  on  this  subject  have  been  g-iven  by  Messrs* 
Murie  and  Mivart,  in  'Transact.  Zoolog.  Soc'  1869,  vol.  vii.  p.  92. 

2*  'Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
pp.  317  and  397.    See  also  'Orig-in  of  Species,'  5th  edit.  p.  535. 

25  For  instance  M.  Richard  ('Annales  des  Sciences  Nat.'  3rd  series, 
Zoolog.  1852,  torn,  xviii.  p.  13)  describes  and  figures  rudiments  of  what 
he  calls  the  "muscle  pedieux  de  la  main,"  which  he  says  is  sometimes 
"infinlment  petit."  Another  muscle  called  "le  tibial  posterieur,"  is 
generally  quite  absent  in  the  hand,  but  appears  from  time  to  time 
in  a  more  or  less  rudimentary  condition. 


RUDIMENTS.  13 

larly  present  in  some  of  the  lower  animals  can  occasionally  be 
detected  in  man  in  a  greatly  reduced  condition.  Every  one  must 
have  noticed  the  power  which  many  animals,  especially  horses, 
possess  of  moving  or  twitching  their  skin;  and  this  is  effected  by 
the  panniculus  carnosus.  Remnants  of  this  muscle  in  an  efficient 
state  are  found  in  various  parts  of  our  bodies;  for  instance,  the 
muscle  on  the  forehead,  by  which  the  eyebrows  are  raised.  The 
platysma  myoides,  which  is  well  developed  on  the  neck,  belongs 
to  this  system.  Prof.  Turner,  of  Edinburgh,  has  occasionally  de- 
tected, as  he  informs  me,  muscular  fasciculi  in  five  different  situ- 
ations, namely  in  the  axillae,  near  the  scapulae,  &c.,  all  of  which 
must  be  referred  to  the  system  of  the  panniculus.  He  has  also 
shown^^  that  the  musculous  sternalis  or  sternalis  brutorum,  which 
is  not  an  extension  of  the  rectus  abdominalis,  but  is  closely  allied 
to  the  panniculus,  occurred  in  the  proportion  of  about  three  per 
cent,  in  upwards  of  600  bodies:  he  adds,  that  this  muscle  affords 
"an  excellent  illustration  of  the  statement  that  occasional  and 
"rudimentary  structures  are  especially  liable  to  variation  in  ar- 
"rangement." 

Some  few  persons  have  the  power  of  contracting  the  super- 
ficial muscles  on  their  scalps;  and  these  muscles  are  in  a  variable 
and  partially  rudimentary  condition.  M.  A.  de  Candolle  has  com- 
municated to  me  a  curious  instance  of  the  long-continued  per- 
sistence or  inheritance  of  this  power,  as  well  as  of  its  unusual  de- 
velopment. He  knows  a  family,  in  which  one  member,  the  pres- 
ent head  of  the  family,  could,  when  a  youth,  pitch  several  heavy 
books  from  his  head  by  the  movement  of  the  scalp  alone;  and  he 
won  wagers  by  performing  this  feat.  His  father,  uncle,  grand- 
father, and  his  three  children  possess  the  same  power  to  the 
same  unusual  degree.  This  family  became  divided  eight  genera- 
tions ago  into  two  branches;  so  that  the  head  of  the  above-men- 
tioned branch  is  cousin  in  the  seventh  degree  to  the  head  of 
the  other  branch.  This  distant  cousin  resides  in  another  part 
of  France;  and  on  being  asked  whether  he  possessed  the  same 
faculty,  immediately  exhibited  his  power.  This  case  offers  a  good 
illustration  how  persistent  may  be  the  transmission  of  an  abso- 
lutely useless  faculty,  probably  derived  from  our  remote  semi- 
human  progenitors;  since  many  monkeys  have,  and  frequently 
use  the  power,  or  largely  moving  their  scalps  up  and  down.^^ 

The  extrinsic  muscles  which  serve  to  move  the  external  ear, 
and  the  intrinsic  muscles  which  move  the  different  parts,  are  in  a 
rudimentary  condition  in  man,  and  they  all  belong  to  the  system 
of  the  panniculus;  they  are  also  variable  in  development,  or  at 


2«  Prof.  W.  Turner,  'Proc.  Royal  Soc.  Edinburg-h,'  1866-67,  p.  65. 
^See  my  'Expression  of  the  Emotions  in  Man  and  Animals,'  1872, 
p.  144. 


14  THE  DESCENT  OF  MAN. 

least  in  function.  I  have  seen  one  man  who  could  draw  the  whole 
ear  forwards;  other  men  can  draw  it  upwards;  an- 
other who  could  draw  it  backwards;-**  and  from  what 
one  of  these  persons  told  me,  it  is  probable  that  most  of  us,  by 
often  touching  our  ears,  and  thus  directing  our  attention  towards 
them,  could  recover  some  power  of  movement  by  repeated  trials. 
The  power  of  erecting  and  directing  the  shell  of  the  ears  to  the 
various  points  of  the  compass,  is  no  doubt  of  the  highest  service 
to  many  animals,  as  they  thus  perceive  the  direction  of  danger; 
but  I  have  never  heard,  on  sufficient  evidence,  of  a  man  who  pos- 
sessed this  power,  the  one  which  might  be  of  use  to  him.  The 
whole  external  shell  may  be  considered  a  rudiment,  together  with 
the  various  folds  and  prominences  (helix  and  anti-helix,  tragus 
and  anti-tragus,  &c.)  which  in  the  lower  animals  strengthen 
and  support  the  ear  when  erect,  without  adding  much  to  its 
weight.  Some  authors,  however,  suppose  that  the  cartilage  of  the 
shell  serves  to  transmit  vibrations  to  the  acoustic  nerve;  but  Mr. 
Toynbee,-''  after  collecting  all  the  known  evidence  on  this  head, 
concludes  that  the  external  shell  is  of  no  distinct  use.  The  ears 
of  the  chimpanzee  and  orang  are  curiously  like  those  of  man, 
and  the  proper  muscles  are  likewise  but  very  slightly  developed.^" 
I  am  also  assured  by  the  keepers  in  the  Zoological  Gardens  that 
these  animals  never  move  or  erect  their  ears;  so  that  they  are 
in  an  equally  rudimentary  condition  with  those  of  man,  as  far  as 
function  is  concerned.  Why  these  animals,  as  well  as  the  pro- 
genitors of  man,  should  have  lost  the  power  of  erecting  their 
ears,  we  cannot  say.  It  may  be,  though  I  am  not  satisfied  with 
this  view,  that  owing  to  their  arboreal  habits  and  great  strength 
they  were  but  little  exposed  to  danger,  and  so  during  a  lengthened 
period  moved  their  ears  but  little,  and  thus  gradually  lost  the 
power  of  moving  them.  This  would  be  a  parallel  case  with  that 
of  those  large  and  heavy  birds,  which,  from  inhabiting  oceanic 
islands,  have  not  been  exposed  to  the  attacks  of  beasts  of  prey, 
and  have  consequently  lost  the  power  of  using  their  wings  for 
flight.  The  inability  to  move  the  ears  in  man  and  several  apes 
is,  however,  partly  compensated  by  the  freedom  with  which  they 
can  move  the  head  in  a  horizontal  plane,  so  as  to  catch  sounds 
from  all  directions.    It  has  been  asserted  that  the  ear  of  man  alone 


28  Canestrini  quotes  Hyrtl.  ('Annuarie  della  Soc.  dei  Naturalist!,'  Mo- 
dena,  1867,  p.  97)  to  the  same  effect. 

29  'The  Diseases  of  the  Ear,'  by  J.  Toynbee,  F.  R.  S.,  1860,  p.  12.  A 
disting-uished  physiologist,  Prof.  Preyer,  informs  me  that  he  had  lately 
been  experimenting'  on  the  function  of  the  shell  of  the  ear,  and  has 
come  to  nearly  the  same  conclusion  as  that  given  here. 

30  Prof.  A.  Macalister,  'Annals  and  Mag-,  of  Nat.  History,'  vol.  vii. 
1871,  p.  342. 


RUDIMENTS. 


15 


possesses  a  lobule;  but  "a  rudiment  of  it  is  found  in  the  gorilla;"^ 
and,  as  I  hear  from  Prof.  Preyer,  it  is  not  rarely  absent  in  the 

negro. 

The  celebrated  sculptor,  Mr.  Woolner,  informs  me  of  one  little 
peculiarity  in  the  external  ear,  which  he  has  often  observed  both 
in  men  and  women,  and  of  which  he  perceived  the  full  signifi- 
cance. His  attention  was  first  called  to  the  subject  whilst  at 
work  on  his  figure  of  Puck,  to  which  he  had  given  pointed  ears. 
He  was  thus  led  to  examine  the  ears  of  various  monkeys,  and  sub- 
sequently more  carefully  those  of  man.  The  peculiarity  consists 
in  a  little  blunt  point,  projecting  from  the  inwardly  folded  margin, 
or  helix.  When  present,  it  is  developed  at  birth,  and,  according 
to  Prof.  Ludwig  Meyer,  more  frequently  in  man  than  in  v/oman. 
Mr.  Woolner  made  an  exact  model  of  one  such  case,  and  sent  me 
the  accompanying  drawing.  (Fig.  2.) 
These  points  not  only  project  inwards 
towards  the  center  of  the  ear,  but  often 
a  little  outwards  from  its  plane,  so  as 
to  be  visible  when  the  head  is  viewed 
from  directly  in  front  or  behind.  They 
are  variable  in  size,  and  somewhat  in 
position,  standing  either  a  little  higher 
or  lower;  and  they  sometimes  occur 
on  one  ear  and  not  on  the  other.  They 
are  not  confined  to  mankind,  for  I  ob- 
served a  case  in  one  of  the  spider- 
monkeys     (Ateles     beelzebuth)     in     our 

Zoological    Gardens;     and    Dr.    E.   Ray     -c.-        o      ^  -p 

Lankester  informs  me  of  another  case  modelled  and'dr awn  by 
in  a  chimpanzee  in  the  gardens  at  Mr.  Woolner. 
Hamburg.  The  helix  obviously  con-  a.  The  projecting  point, 
sists  of  the  extreme  margin  of  the  ear  folded  inwards;  and  this 
folding  appears  to  be  in  some  manner  connected  with  the  whole 
external  ear  being  permanently  pressed  backwards.  In  many 
monkeys,  which  do  not  stand  high  in  the  order,  as  baboons  and 
some  species  of  macacus,^^  the  upper  portion  of  the  ear  is  slightly 
pointed,  and  the  margin  is  not  at  all  folded  inwards;  but  if  the 
margin  were  to  be  thus  folded,  a  slight  point  would  necessarily 
project  inwards  towards  the  center,  and  probably  a  little  out- 
wards from  the  plane  of  the  ear;  and  this  I  believe  to  be  their 
origin  in  many  cases.  On  the  other  hand.  Prof.  L.  Meyer,  in  an 
able  paper  recently  published,^^  maintains  that  the  whole  case  is 


31  Mr.  St.  George  Mivart,  'Elementary  Anatomy,'  1873,  p.  396. 

32  See  also  some  remarks,  and  the  drawings  of  the  ears  of  the  Lemu- 
roidea,  in  Messrs.  Murie  and  Mivart's  excellent  paper  in  'Transact. 
Zoolog.  Soc'  vol.  vii.  1869,  pp.  6  and  90. 

33  Ueber  das  Darwin'sche  Spitzohr,  Archiv  fur  Path.  Anat.  und  Phys. 
1871,  p.  485. 


16  THE  DESCENT  OP  MAN. 

one  of  mere  variability;  and  that  the  projections  are  not  real 
ones,  but  are  due  to  the  internal  cartilage  on  each  side  of  the 
points  not  having  been  fully  developed.  I  am  quite  ready  to 
admit  that  this  is  the  correct  explanation  in  many  instances,  as 
in  those  figured  by  Prof.  Meyer,  in  which  there  are  several  minute 
points,  or  the  whole  margin  is  sinuous.  I  have  myself  seen, 
through  the  kindness  of  Dr.  L.  Down,  the  ear  of  a  microcephalous 
idiot,  on  which  there  is  a  projection  on  the  outside  of  the  helix, 
and  not  on  the  inward  folded  edge,  so  that  this  point  can  have 
no  relation  to  a  former  apex  of  the  ear.  Nevertheless  in  some 
cases,  my  original  view,  that  the  points  are  vestiges  of  the  tips 
of  formerly  erect  and  pointed  ears,  still  seems  to  me  probable.  I 
think  so  from  the  frequency  of  their  occurrence,  and  from  the  gen- 
eral correspondence  in  position  with  that  of  the  tip  of  a  pointed 
ear.  In  one  case,  of  which  a  photograph  has  been  sent  me,  the 
projection  is  so  large,  that  supposing,  in  accordance  with  Prof. 
Meyer's  view,  the  ear  to  be  made  perfect  by  the  equal  development 
of  the  cartilage  throughout  the  whole  extent  of  the  margin,  it 
would  have  covered  fully  one-third  of  the  whole  ear.  Two  cases 
have  been  communicated  to  me,  one  in  North  America,  and  the 
other  in  England,  in  which  the  upper  margin  is  not  at  all  folded 
inwards,  but  is  pointed,  so  that  it  closely  resembles  the  pointed 
ear  of  an  ordinary  quadruped  in  outline.  In  one  of  these  cases, 
which  was  that  of  a  young  child,  the  father  compared  the  ear  with 
the  drawing  which  I  have  given^*  of  the  ear  of  a  monkey,  the 
Cynopithecus  niger,  and  says  that  their  outlines  are  closely  sim- 
ilar. If,  in  these  two  cases,  the  margin  had  been  folded  inwards 
in  the  normal  manner,  an  inward  projection  must  have  been 
formed.  I  may  add  that  in  two  other  cases  the  outline  still  re- 
mains somewhat  pointed,  although  the  margin  of  the  upper  part 
of  the  ear  is  normally  folded  inwards — in  one  of  them,  however, 
very  narrowly.  The  follow^ing  woodcut  (No.  3)  is  an  accurate 
copy  of  a  photograph  of  the  foetus  of  an  orang  (kindly  sent  me 
by  Dr.  Nitsche),  in  which  it  may  be  seen  how  different  the  pointed 
outline  of  the  ear  is  at  this  period  from  its  adult  condition,  when 
it  bears  a  close  general  resemblance  to  that  of  man.  It  is  evident 
that  the  folding  over  of  the  tip  of  such  an  ear,  unless  it  changed 
greatly  during  its  further  development,  would  give  rise  to  a  point 
projecting  inwards.  On  the  whole,  it  still  seems  to  me  probable 
that  the  points  in  question  are  in  some  cases,  both  in  man  and 
apes,  vestiges  of  a  former  condition. 

The  nictitating  membrane,  or  third  eyelid,  with  its  accessory 
muscles  and  other  structures,  is  especially  well  developed  in 
birds,  and  is  of  much  functional  importance  to  them  as  it  can 
be  rapidly  drawn  across  the  whole  eye-ball.    It  is  found  in  some 

8*  'The  Expression  of  the  Emotions,'  p.  136. 


HtJDIMENTS.  17 

reptiles  and  amphibians,  and  in  certain  fishes,  as  in  sharks.  It 
is  fairly  well  developed  in  the  two  lower  divisions  of  the  mam- 
malian series,  namely,  in  the  monotremata  and  marsupials,  and 
in  some  few  of  the  higher  mammals,  as  in  the  walrus.    But  in 


Fig.  3.    Foetus  of  an  Orang.    Exact  copy  of  a  photograph,  showing 
the  form   of  the   ear  at  this   early   age. 

man,  the  quadrumana,  and  most  other  mammals,  it  exists,  as  is 
admitted  by  all  anatomists,  as  a  mere  rudiment,  called  the 
semilunar  fold.^^ 

The  sense  of  smell  is  of  the  highest  importance  to  the  greater 
number  of  mammals — to  some,  as  the  ruminants,  in  warning 
them  of  danger;  to  others,  as  the  carnivora,  in  finding  their  prey; 
to  others,  again,  as  the  wild  boar,  for  both  purposes  combined. 
But  the  sense  of  smell  is  of  extremely  slight  service,  if  any,  even 
to  the  dark  colored  races  of  men,  in  whom  it  is  much  more  highly 
developed  than  in  the  white  and  civilized  races.^^  Nevertheless  it 

85  Muller's  'Elements  of  Physiology,'  Eng.  translat.,  1842,  vol,  ii.  p. 
1117.  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  260;  ibid  on  the  Wal- 
rus, 'Proc.  Zoolog.  Soc'  November  8th,  1854.  See  also  R.  Knox,  'Great 
Artists  and  Anatomists,'  p.  106.  This  rudiment  apparently  is  somewhat 
larger  in  Negroes  and  Australians  than  in  Europeans,  see  Carl  Vogt, 
'Lectures  on  Man,'  Eng.  trajislat.  p.  129. 

3«  The  account  given  by  Humboldt  of  the  power  of  smell  possessed  by 
the  natives  of  South  America  is  well  known,  and  has  been  confirmed 
by  others.  M.  Houzeau  ('Etudes  sur  les  Facultes  Mentales,'  &c.,  torn. 
i.  1872,  p.  91)  asserts  that  he  repeatedly  made  experiments,  and  proved 
that  Negroes  and  Indians  could  recognize  persons  in  the  dark  by  their 
odor.  Dr.  W.  Ogle  has  made  some  curious  observations  on  the  con- 
nection between  the  power  of  smell  and  the  coloring  matter  of  the  mu- 


18  THE  DESCENT  OF  MAN. 

does  not  warn  them  of  danger,  nor  guide  them  to  their  food;  nor 
does  it  prevent  the  Esquimaux  from  sleeping  in  the  most  fetid  at- 
mosphere, nor  many  savages  from  eating  half-putrid  meat.  In 
Europeans  the  power  differs  greatly  in  different  individuals,  as  I 
am  assured  by  an  eminent  naturalist  who  possesses  this  sense 
highly  developed,  and  who  has  attended  to  the  subject.  Those 
who  believe  in  the  principle  of  gradual  evolution,  will  not  readily 
admit  that  the  sense  of  smell  in  its  present  state  was  originally 
acquired  by  man,  as  he  now  exists.  He  inherits  the  power  in  an 
enfeebled  and  so  far  rudimentary  condition,  from  some  early  pro- 
genitor, to  whom  it  was  highly  serviceable,  and  by  whom  it  was 
continually  used.  In  those  animals  which  have  this  sense  highly 
developed,  such  as  dogs  and  horses,  the  recollection  of  persons 
and  of  places  is  strongly  associated  with  their  odor;  and  we  can 
thus  perhaps  understand  how  it  is,  as  Dr.  Maudsley  has  truly  re- 
marked," that  the  sense  of  smell  in  man  "is  singularly  effective 
"in  recalling  vividly  the  ideas  and  images  of  forgotten  scenes  and 
"places." 

Man  differs  conspicuously  from  all  tihe  other  Primates  in  being 
almost  naked.  But  a  few  short  straggling  hairs  are  found  over 
the  greater  part  of  the  body  in  the  man,  and  fine  down  on  that  of 
the  woman.  The  different  races  differ  much  in  hairiness;  and  in 
the  individuals  of  the  same  race  the  hairs  are  highly  variable, 
not  only  in  abundance,  but  likewise  in  position:  thus  in  some 
Europeans  the  shoulders  are  quite  naked,  whilst  in  others  they 
bear  thick  tufts  of  hair.^«  There  can  be  little  doubt  that  the  hairs 
thus  scattered  over  the  body  are  the  rudiments  of  the  uniform 
hairy  coat  of  the  lower  animals.  This  view  is  rendered  all  the 
more  probable,  as  it  is  known  that  fine,  short,  and  pale-colored 
hairs  on  the  limbs  and  other  parts  of  the  body,  occasionally  be- 
come developed  into  "thickset,  long,  and  rather  coarse  dark 
"hairs,"  when  abnormally  nourished  near  old-standing  infiamed 
surfaces.^®  / 

I  am  informed  by  Sir  James  Paget  that  often  several  members 
of  a  family  have  a  few  hairs  in  their  eyebrows  much  longer  than 
the  others;  so  that  even  this  slight  peculiarity  seems  to  be  in- 
herited. These  hairs,  too,  seem  to  have  their  representatives;  for 
in  the  chimpanzee,  and  in  certain  species  of  Macacus,  there  are 


cous  membrane  of  the  olfactory  region,  as  well  as  of  the  skin  of  the 
body.    I  have,  therefore,  spoken  in  the  text  of  the  dark-colored  races 
having-  a  finer  sense  of  smell  than  the  white  races.    See  his  paper  'Med- 
ico-Chirurgical  Transactions,'  London,  vol.  liii.,  1870,  p.  276. 
87  'The  Physiology  and  Pathology  of  Mind,'  2nd  edit.  1868,  p.  134. 

38  Eschricht,  Ueber  die  Richtung  der  Haare  am  menschlichen  Korper, 
'Muller's  Archiv  fur  Anat.  und  Phys.'  1837,  s.  47.  I  shall  often  have 
to  refer  to  this  very  curious  paper. 

39  Paget,  'Lectures  on  Surgical  Pathology,'  1853,  vol.  i.  p.  71. 


RUDIMENTS.  19 

scattered  hairs  of  considerable  length  rising  from  the  naked  skin 
above  the  eyes,  and  corresponding  to  our  eyebrows;  similar  long 
hairs  project  from  the  hairy  covering  of  the  superciliary  ridges  in 
some  baboons. 

The  fine  wool-like  hair,  or  so-called  lanugo,  with  which  the 
human  foetus  during  the  sixth  month  is  thickly  covered,  offers  a 
more  curious  case.  It  is  first  developed,  during  the  fifth  month, 
on  the  eyebrows  and  face,  and  especially  round  the  mouth,  where 
it  is  much  longer  than  that  on  the  head.  A  mustache  of  this 
kind  was  observed  by  Eschricht^"  on  a  female  foetus;  but  this  is 
not  so  surprising  a  circumstance  as  it  may  first  appear,  for  the 
two  sexes  generally  resemble  each  other  in  all  external  characters 
during  an  early  period  of  growth.  The  direction  and  arrange- 
ment of  the  hairs  on  all  parts  of  the  foetal  body  are  the  same 
as  in  the  adult,  but  are  subject  to  much  variability.  The  whole 
surface,  including  even  the  forehead  and  ears,  is  thus  thickly 
clothed;  but  it  is  a  significant  fact  that  the  palms  of  the  hands 
and  the  soles  of  the  feet  are  quite  naked,  like  the  inferior  surfaces 
of  all  four  extremities  in  most  of  the  lower  animals.  As  this 
can  hardly  be  an  accidental  coincidence,  the  woolly  covering  of 
the  foetus  probably  represents  the  first  permanent  coat  of  hair 
in  those  mammals  which  are  born  hairy.  Three  or  four  cases  have 
been  recorded  of  persons  born  with  their  whole  bodies  and  faces 
thickly  covered  with  fine  long  hairs;  and  this  strange  condition 
is  strongly  inherited,  and  is  correlated  with  an  abnormal  condition 
of  the  teeth.*^  Prof.  Alex.  Brandt  informs  me  that  he  has  com- 
pared the  hair  from  the  face  of  a  man  thus  characterized,  aged 
thirty-five,  with  the  lanugo  of  a  foetus,  and  finds  it  quite  similar 
in  texture;  therefore,  as  he  remarks,  the  case  may  be  attributed 
to  an  arrest  of  development  in  the  hair,  together  with  its  con- 
tinued growth.  Many  delicate  children,  as  I  have  been  assured 
by  a  surgeon  to  a  hospital  for  children,  have  their  backs  covered 
by  rather  long  silky  hairs;  and  such  cases  probably  come  under 
the  same  head. 

It  appears  as  if  the  posterior  molar  or  wisdom-teeth  were  tend- 
ing to  become  rudimentary  in  the  more  civilized  races  of  man. 
These  teeth  are  rather  smaller  than  the  other  molars,  as  is  like- 
wise the  case  with  the  corresponding  teeth  in  the  chimpanzee  and 
orang;  and  they  have  only  two  separate  fangs.  They  do  not  cut 
through  the  gums  till  about  the  seventeenth  year,  and  I  have  been 
assured  that  they  are  much  more  liable  to  decay,  and  are  earlier 
lost  than  the  other  teeth;    but  this  is  denied  by  some  eminent 


*o  Eschricht,  ibid.  s.  40,  47. 

*^  See  my  'Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
ii.  p.  327.  Prof.  Alex.  Brandt  has  recently  sent  me  an  additional  case 
of  a  father  and  son,  born  in  Russia,  with  thes.e  peiculiaritie&.  I  have 
received  drawing-s  of  both  from  Paris. 


20  THE  DESCENT  OF  MAN. 

dentists.  They  are  also  much  more  liable  to  vary,  both  in  structure 
and  in  the  period  of  their  development,  than  the  other  teeth.^^  j^ 
the  Melanian  races,  on  the  other  hand,  the  wisdom-teeth  are  usual- 
ly furnished  with  three  separate  fangs,  and  are  generally  sound; 
they  also  differ  from  the  other  molars  in  size,  less  than  in  the  Cau- 
casian race.*8  Prof.  Schaaffhausen  accounts  for  this  difference  be- 
tween the  races  by  "the  posterior  dental  portion  of  the  jaw  being 
"always  shortened"  in  those  that  are  civilized,**  and  this  shorten- 
ing may,  I  presume,  be  attributed  to  civilized  men  habitually 
feeding  on  soft,  cooked  food,  and  thus  using  their  jaws  less.  I  am 
informed  by  Mr.  Brace  that  it  is  becoming  quite  a  common  prac- 
tice in  the  United  States  to  remove  some  of  the  molar  teeth  of 
children,  as  the  jaw  does  not  grow  large  enough  for  the  perfect 
development  of  the  normal  number.*^ 

With  respect  to  the  alimentary  canal,  I  have  met  with  an  ac- 
count of  only  a  single  rudiment,  namely  the  vermiform  appendage 
of  the  caecum.  The  caecum  is  a  branch  or  diverticulum  of  the  intes- 
tine, ending  in  a  cul-de-sac,  and  is  extremely  long  in  many  of  the 
lower  vegetable-feeding  mammals.  In  the  marsupial  koala  it  is  ac- 
tually more  than  thrice  as  long  as  the  whole  body.*''  It  is  some- 
times produced  into  a  long  gradually-tapering  point,  and  is  some- 
times constricted  in  parts.  It  appears  as  if,  in  consequence  of 
changed  diet  or  habits,  the  caecum  had  become  much  shortened  in 
various  animals,  the  vermiform  appendage  being  left  as  a  rudi- 
ment of  the  shortened  part.  That  this  appendage  is  a  rudiment,  we 
may  infer  from  its  small  size,  and  from  the  evidence  which  Prof. 
Canestrini*^  has  collected  of  its  variability  in  man.  It  is  occasion- 
ally quite  absent,  or  again  is  largely  developed.  The  passage  is 
sometimes  completely  closed  for  half  or  two-thirds  of  its  length, 
with  the  terminal  part  consisting  of  a  flattened  solid  expansion.  In 
the  orang  this  appendage  is  long  and  convoluted:  in  man  it  arises 
from  the  end  of  the  short  caecum,  and  is  commonly  from  four  to 
five  inches  in  length,  being  only  about  the  third  of  an  inch  in 
diameter.  Not  only  is  it  useless,  but  it  is  sometimes  the  cause  of 
death,  of  which  fact  I  have  lately  heard  two  instances:  this  is  due 
to  small  hard  bodies,  such  as  seeds,  entering  the  passage,  and 
causing  inflammation.*^ 

*2  Dr.  Webb,  'Teeth  in  Man  and  the  Anthropoid  Apes,'  as  quoted  by 
Dr.  C.  Carter  Blake  in  'Anthropological  Review,'  July,  1867,  p.  299. 

^  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  pp.  320,  321  and  325. 

**  'On  the  Primitive  Form  of  the  Skull,'  Eng-.  translat.  in  'Anthropo- 
logical Review,'  Oct.  1868,  p.  426. 

^  Prof.  Mantegazza  writes  to  me  from  Florence  that  he  has  lately 
been  studying  the  last  molar  teeth  in  the  different  races  of  man,  and 
has  come  to  the  same  conclusion  as  that  given  in  my  text,  viz.,  that 
in  the  higher  or  civilized  races  they  are  on  the  road  towards  atrophy 
or  elimination. 

*«  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  pp.  416,  434,  441. 

♦^  'Annuario   della   Soc.   d.   Nat.'   Modena,   1867,   p.   94. 

*»M.  C.  Martins  ("De  I'Unite  Organique,"  in  'Revue  des  Deux  Mon- 


RUDIMEN-rS.  21 

In  St5me  of  the  lower  Quadrumana,  in  the  Lemuridas  and  Carni- 
vora,  as  well  as  in  many  marsupials,  there  is  a  passage  near  the 
loYv^er  end  of  the  humerus,  called  the  supra-condyloid  foramen, 
through  which  the  great  nerve  of  the  fore  limb  and  often  the  great 
artery  pass.  Now  in  the  humerus  of  man,  there  is  generally  a  trace 
of  this  passage,  which  i's  sometimes  fairly  well  developed,  being 
formed  by  a  depending  hook-like  process  of  bone,  completed  by  a 
band  of  ligament.  Dr.  Struthers,^''  who  has  closely  attended  to  the 
subject,  has  now  shown  that  this  peculiarity  is  sometimes  inherit- 
ed, as  it  has  occurred  in  a  father,  and  in  no  less  than  four  out  of 
his  seven  children.  When  present,  the  great  nerve  invariably 
passes  through  it;  and  this  clearly  indicates  that  it  is  the  homo- 
logue  and  rudiment  of  the  «upra-condyloid  foramen  of  the  lower 
animals.  Prof.  Turner  estimates,  as  he  informs  me,  that  it  oc- 
curs in  about  one  per  cent,  of  recent  skeletons.  But  if  the  occa- 
sional development  of  this  structure  in  man  is,  as  seems  probable, 
due  to  reversion,  it  is  a  return  to  a  very  ancient  state  of  things,  be- 
cause in  the  higher  Quadrumana  it  is  absent. 

There  is  another  foramen  or  perforation  in  the  humerus,  occa- 
sionally present  in  man,  which  may  be  called  the  inter-condyioid. 
This  occurs,  but  not  constantly,  in  various  anthropoid  and  other 
apes,^"  and  likewise  in  many  of  the  lower  animals.  It  is  remark- 
able that  this  perforation  seems  to  have  been  present  in  man  much 
more  frequently  during  ancient  times  than  recently.  Mr.  Busk^^ 
has  collected  the  following  evidence  on  this  head:  Prof.  Broca  "no- 
"ticed  the  perforation  in  four  and  a  half  per  cent,  of  the  arm-bones 
"collected  in  the  'Cimetiere  du  Sud,'  at  Paris;  and  in  the  Grotto  of 
"Orrony,  the  contents  of  which  are  referred  to  the  Bronze  period, 
"as  many  as  eight  humeri  out  of  thirty-two  were  perforated;  but 
"this  extraordinary  proportion,  he  thinks,  might  be  due  to  the  cav- 
"ern  having  been  a  sort  of  'family  vault,'  Again,  M.  Dupont  found 
"thirty  per  cent,  of  perforated  bones  in  the  caves  of  the  Valley  of 
"the  Lesse,  belonging  to  the  Reindeer  period;  whilst  M.  Leguay,  in 

des,'  June  15,  1862,  p.  16),  and  Hackel  ('Generelle  Morphologie,'  B.  ii.  s. 
278),  have  both  remarked  on  the  singular  fact  of  this  rudiment  some- 
times causing-  death. 

*9  With  respect  to  inheritance,  see  Dr.  Struthers  in  the  'Lancet,'  Feb. 
15,  1873,  and  another  important  paper,  ibid.,  Jan.  24,  1863,  p.  83.  Dr. 
Knox,  as  I  am  informed,  was  the  first  anatomist  who  drew  attention 
to  this  peculiar  structure  in  man;  see  his  'Great  Artists  and  Anato- 
mists,' p.  63.  See  also  an  important  memoir  on  this  process  by  Dr. 
Gruber,  in  the  'Bulletin  de  I'Acad.  Imp.  de  St.  Petersbourg,  torn,  xii 
1867,  p.  448. 

50  Mr.  St.  George  Mivart,  'Transact.  Phil.  Soc'  1867,  p.  310. 

51  "On  the  Caves  of  Gibraltar,"  'Transact.  Internat.  Congress  of  Pre- 
hlst.  Arch.'  Third  Session,  18G9,  p.  159.  Prof.  Vv^yman  has  lately  shown 
(Fourth  Annual  Report,  Peabody  Museum,  1871,  p.  20),  that  this  per- 
foration is  present  in  thirty-one  per  cent,  of  some  human  remains 
from  ancient  mounds  in  the  Western  United  States,  and  in  Florida. 
It  frequently  occurs  in  the  negro. 

3 


2^  THE  DESCENT  OP  MAN. 

"a  sort  of  dolmen  at  Argenteuil,  observed  twenty-five  per  cent,  to 
"be  perforated;  and  M.  Pruner-Bey  found  twenty-six  per  cent,  in 
"the  same  condition  in  bones  from  Vaureal.  Nor  should  it  be  left 
"unnoticed  that  M.  Pruner-Bey  states  that  this  condition  is  com- 
■'mon  in  Guanche  skeletons."  It  is  an  interesting  fact  that  ancient 
races,  in  this  and  several  other  cases,  more  frequently  present 
structures  which  resemble  those  of  the  lower  animals  than  do  the 
modern.  One  chief  cause  seems  to  be  that  the  ancient  races  stand 
somewhat  nearer  in  the  long  line  of  descent  to  their  remote  ani- 
mal-like progenitors. 

In  man,  the  os  coccyx,  together  with  certain  other  vertebrae  here- 
after to  be  described,  though  functionless  as  a  tail,  plainly  repre- 
sent this  part  in  other  vertebrate  animals.  At  an  early  embryonic 
period  it  is  free,  and  projects  beyond  the  lower  extremities;  as  may 
be  seen  in  the  drawing  (Fig.  1.)  of  a  human  embryo.  Even  after 
birth  it  has  been  known,  in  certain  rare  and  anomalous  cases,^^  to 
form  a  small  external  rudiment  of  a  tail.  The  os  coccyx  is  short, 
usually  including  only  four  vertebrse,  all  anchylosed  together:  and 
these  are  in  a  rudimentary  condition,  for  they  consist,  with  the 
exception  of  the  basal  one,  of  the  centrum  alone.^^  They  are  fur- 
nished with  some  small  muscles;  one  of  which,  as  I  am  informed 
by  Prof.  Turner,  has  been  expressly  described  by  Theile  as  a  ru- 
dimentary repetition  of  the  extensor  of  the  tail,  a  muscle  which 
is  so  largely  developed  in  many  mammals. 

The  spinal  cord  in  man  extends  only  as  far  downwards  as  the 
last  dorsal  or  first  lumbar  vertebra;  but  a  thread-like  structure 
(the  filum  terminale)  runs  down  the  axis  of  the  sacral  part  of  the 
spinal  canal,  and  even  along  the  back  of  the  coccygeal  bones.  The 
upper  part  of  this  filament,  as  Prof.  Turner  informs  me,  is  un- 
doubtedly homologous  with  the  spinal  cord;  but  the  lower  part  ap- 
parently consists  merely  of  the  pia  mater,  or  vascular  investing 
membrane.  Even  in  this  case  the  os  coccyx  may  be  said  to  possess 
a  vestige  of  so  important  a  structure  as  the  spinal  cord,  though  no 
longer  enclosed  within  a  bony  canal.  The  following  fact,  for  which 
I  am  also  indebted  to  Prof.  Turner,  shows  how  closely  the  os  coc- 
cyx corresponds  with  the  true  tail  in  the  lower  animals:  Luschka 
has  recently  discovered  at  the  extremity  of  the  coccygeal  bones  a 
very  peculiar  convoluted  body,  which  is  continuous  with  the  mid- 
dle sacral  artery;  and  this  discovery  led  Krause  and  Meyer  to  ex- 
amine the  tail  of  a  monkey  (Macacus),  and  of  a  cat,  in  both  of 

S2  Quatrefages  has  lately  collected  the  evidence  on  this  subject. 
'Revue  des  Cours  Scientifiques,'  1867-1868,  p.  625.  In  1840  Fleischmann 
exhibited  a  human  foetus  bearing  a  free  tail,  which,  as  is  not  always 
the  case,  included  vertebral  bodies;  and  this  tail  was  critically  exam- 
ined by  the  many  anatomists  present  at  the  meeting  of  naturalists  at 
Erlang-en  (see  Marshall  in  Niederlandischen  Archiv  fur  Zoologie,  Pp* 
cember,  1871.) 

63  Owen,  'On  the  Nature  of  Limbs,'  1849,  p.  114. 


RUDIMENTS.  23 

■whicli  they  found  a  similarly  convoluted  body,  though  not  at  the 
extremity. 

The  reproductive  system  offers  various  rudimentary  structures; 
but  these  differ  in  one  important  respect  from  the  foregoing 
cases.  Here  we  are  not  concerned  with  the  vestige  of  a  part  which 
does  not  belong  to  the  species  in  an  efficient  state,  but  with  a 
part  efficient  in  the  one  sex,  and  represented  in  the  other  by  a 
mere  rudiment.  Nevertheless,  the  occurrence  of  such  rudiments 
is  as  difficult  to  explain,  on  the  belief  of  the  separate  creation  of 
each  species,  as  in  the  foregoing  cases.  Hereafter  I  shall  have  to 
recur  to  these  rudiments,  and  shall  show  that  their  presence  gen- 
erally depends  merely  on  inheritance,  that  is,  on  parts  acquired 
by  one  sex  having  been  partially  transmitted  to  the  other.  I  will 
in  this  place  only  give  some  instances  of  such  rudiments.  It  is 
well  known  that  in  the  males  of  all  mammals,  including  man,  rudi- 
mentary mammae  exist.  These  in  several  instances  have  become 
well  developed,  and  have  yielded  a  copious  supply  of  milk.  Their 
essential  identity  in  the  two  sexes  is  likewise  shown  by  their  oc- 
casional sympathetic  enlargement  in  both  during  an  attack  of  the 
measles.  The  vesicula  prostatica,  which  has  been  observed  in 
many  male  mammals,  is  now  universally  acknowledged  to  be 
the  homologue  of  the  female  uterus,  together  with  the  connected 
passage.  It  is  impossible  to  read  Leuckart's  able  description 
of  this  organ,  and  his  reasoning,  without  admitting  the  justness 
of  his  conclusion.  This  is  especially  clear  in.  the  case  of  those 
mammals  in  which  the  true  female  uterus  bifurcates,  for  in  the 
males  of  these  the  vesicula  likewise  bifurcates.^*  Some  other  rudi- 
mentary structures  belonging  to  the  reproductive  system  might 
have  been  here  adduced.^^ 

The  bearing  of  the  three  great  classes  of  facts  now  given  is 
unmistakable.  But  it  would  be  superfluous  fully  to  recapitulate 
the  line  of  argument  given  in  detail  in  my  'Origin  of  Species.' 
The  homological  construction  of  the  whole  frame  in  the  members 
of  the  same  class  is  intelligible,  if  we  admit  their  descent  from 
a  common  progenitor,  together  with  their  subsequent  adaptation 
to  diversified  conditions.  On  any  other  view,  the  similarity  of 
pattern  between  the  hand  of  a  man  or  monkey,  the  foot  of  a 
horse,  the  flipper  of  a  seal,  the  wing  of  a  bat,  &c.,  is  utterly 
inexplicable.^®    It  is  no  scientific  explanation  to  assert  that  they 


B4Leuckart,  in  Todd's  'Cyclop,  of  Anat.'  1849-52,  vol.  iv.  p.  1415.  In 
man  this  organ  is  only  from  three  to  six  lines  in  length,  but,  like  so 
many  other  rudimentary  parts,  it  is  variable  in  development  as  well  as 
in  other  characters. 

^  See,  on  this  subject,  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  pp. 
675,  676,  706. 

^  Prof.  Bianconi,  in  a  recently  published  work,  illustrated  by  ad- 
mirable engravings  ('La  Theorie  Darwinienne  et  la  creation  dite  inde- 
pendante,' .  1874),  endeavors  to  show  that  homological  structures,  in 
the  above  and  other  cases,  can  be  fully  explained  on  mechanical  prin» 


24  THE  DESCENT  OF  MAN. 

have  all  been  formed  on  the  same  ideal  plan.  With  respect  to 
development,  we  can  clearly  understand,  on  the  principle  of  varia- 
tions supervening  at  a  rather  late  embryonic  period,  and  being 
inherited  at  a  corresponding  period,  how  it  is  that  the  embryos 
of  wonderfully  different  forms  should  still  retain,  more  or  less 
perfectly,  the  structure  of  their  common  progenitor.  No  other 
explanation  has  ever  been  given  of  the  marvelous  fact  that  the 
embryos  of  a  man,  dog,  seal,  bat,  reptile,  &c.,  can  at  first  hardly 
be  distinguished  from  each  other.  In  order  to  understand  the 
existence  of  rudimentary  organs,  we  have  only  to  suppose  that  a 
former  progenitor  possessed  the  parts  in  question  in  a  perfect 
state,  and  that  under  changed  habits  of  life  they  became  greatly 
reduced,  either  from  simple  disuse,  or  through  the  natural  selec- 
tion of  those  individuals  which  were  least  encumbered  with  a 
superfluous  part,  aided  by  the  other  means  previously  indicated. 

Thus  we  can  understand  how  it  has  come  to  pass  that  man  and 
all  other  vertebrate  animals  have  been  constructed  on  the  same 
general  model,  why  they  pass  through  the  same  early  stages  of 
development,  and  why  they  retain  certain  rudiments  in  common. 
Consequently  we  ought  frankly  to  admit  their  community  of 
descent;  to  take  any  other  view,  is  to  admit  that  our  own  struc- 
ture, and  that  of  all  the  animals  around  us,  is  a  mere  snare  laid 
to  entrap  our  judgment.  This  conclusion  is  greatly  strengthened, 
if  we  look  to  the  members  of  the  whole  animal  series,  and  con- 
sider the  evidence  derived  from  their  affinities  or  classification, 
their  geographical  distribution  and  geological  succession.  It  is 
only  our  natural  prejudice,  and  that  arrogance  which  made  our 
forefathers  declare  that  they  were  descended  from  demi-gods, 
which  leads  us  to  demur  to  this  conclusion.  But  the  time  will  be- 
fore long  come,  when  it  will  be  thought  wonderful  that  naturalists, 
who  were  well  acquainted  with  the  comparative  structure  and  de- 
velopment of  man,  and  other  mammals,  should  have  believed  that 
each  was  the  work  of  a  separate  act  of  creation. 

ciples  in  accordance  with  their  uses.  No  one  has  shown  so  well,  how 
admirably  such  structures  are  adapted  for  their  final  purpose;  and 
this  adaptation  can,  as  I  believe,  be  explained  through  natural  selection. 
In  considering  the  wing  of  a  bat,  he  brings  forward  (p.  218)  what  ap- 
pears to  me  (to  use  Auguste  Comte's  words)  a  mere  metaphysical 
principle  namely,  the  preservation  "in  its  integrity  of  the  mammalian 
"nature  of  the  animal."  In  only  a  few  cases  does  he  discuss  rudiments, 
and  then  only  those  parts  which  are  partially  rudimentary,  such  as  the 
little  hoofs  of  the  pig  and  ox,  which  do  not  touch  the  ground;  these 
he  shows  clearly  to  be  of  service  to  the  animal.  It  is  unfortunate  that 
he  did  not  consider  such  cases  as  the  minute  teeth,  which  never  cut 
through  the  jaw  in  the  ox,  or  the  mammae  of  male  quadrupeds,  or  the 
wings  of  certain  beetles,  existing  under  the  soldered  wing-covers,  or 
the  vestiges  of  the  pistil  and  stamens  in  various  flov/ers,  and  many 
other  such  cases.  Although  I  greatly  admire  Prof.  Bianconi's  work, 
yet  the  belief  now  held  by  most  naturalists  seems  to  me  left  unshaken, 
that  homological  structures  are  inexplicable  on  the  principle  of  mere 
adaptation. 


MANNER  OF  DEVELOPMENT.  2£ 


CHAPTER   II. 

ON    THE    MANNER    OF    DEVELOPMENT    OF    MAN 
FROM    SOME    LOWER    FORM. 

Variability  of  body  and  mind  in  man— Inheritance— Causes  of  varia- 
bility—Laws of  variation  the  same  in  man  as  in  the  lower  animals- 
Direct  action  of  the  conditions  of  life — Effects  of  the  increased  use 
and  disuse  of  parts— Arrested  development— Reversion— Correlated 
variation— Rate  of  increase — Checks  to  increase — Natural  selection — 
Man  the  most  dominant  animal  in  the  world — Importance  of  his  cor- 
poreal structure — The  causes  which  have  led  to  his  becoming-  erect^ 
Consequent  chang-es  of  structure — Decrease  in  size  of  the  canine 
teeth— Increased  size  and  altered  shape  of  the  skull — ^Nakedness — 
Absence  of  a  tail— Defenseless  condition  of  man. 

It  is  manifest  that  man  is  now  subject  to  much  variability.  No 
two  individuals  of  the  same  race  are  quite  alike.  We  may  compare 
millions  of  faces,  and  each  will  be  distinct.  There  is  an  equally 
great  amount  of  diversity  in  the  proportions  and  dimensions  of 
the  various  parts  of  the  body;  the  length  of  the  legs  being  one 
of  the  most  variable  points.^  Although  in  some  quarters  of  the 
world  an  elongated  skull,  and  in  other  quarters  a  short  skull 
prevails,  yet  there  is  great  diversity  of  shape  even  within  the 
limits  of  the  same  race,  as  with  the  aborigines  of  America  and 
South  Australia — the  latter  a  race  "probably  as  pure  and  homo- 
"geneous  in  blood,  customs,  and  language  as  any  in  existence" — 
and  even  with  the  inhabitants  of  so  confined  an  area  as  the  Sand- 
wich Islands.^  An  eminent  dentist  assures  me  that  there  is  nearly 
as  much  diversity  in  the  teeth  as  in  the  features.  The  chief  ar- 
teries so  frequently  run  in  abnormal  courses,  that  it  has  been 
found  useful  for  surgical  purposes  to  calculate  from  1040  corpses 

1  'Investigations  in  Military  and  Anthropolog.  Statistics  of  American 
Soldiers,'  by  B.  A.  Gould,  1869,  p.  256. 

2  With  respect  to  the  "Cranial  forms  of  the  American  aborigines," 
see  Dr.  Aitken  Meigs  in  'Proc.  Acad.  Nat.  Sci.'  Philadelphia,  May,  1868, 
On  the  Australians  see  Huxley,  in  Lyell's  'Antiquity  of  Man,'  1863,  p. 
87.  On  the  Sandwich  Islanders,  Prof.  J.  Wym^an,  'Observations  on  Cra- 
nia,' Boston,  1868,  p.  18. 


26  THE  DESCENT  OF  MAN. 

how  often  each  course  prevails.^  The  muscles  are  eminently 
variable:  thus  those  of  the  foot  were  found  by  Prof.  Turner^  not 
to  be  strictly  alike  in  any  two  out  of  fifty  bodies;  and  in  some 
the  deviations  were  considerable.  He  adds,  that  the  power  of  per- 
forming the  appropriate  movements  must  have  been  modified  in 
accordance  with  the  several  deviations.  Mr.  J.  Wood  has  recorded' 
the  occurrence  of  295  muscular  variations  in  thirty-six  subjects, 
and  in  another  set  of  the  same  number  no  less  than  558  variations, 
those  occurring  on  both  sides  of  the  body  being  only  reckoned  as 
one.  In  the  last  set,  not  one  body  out  of  the  thirty-six  was 
"found  totally  wanting  in  departures  from  the  standard  descrip- 
"tions  of  the  muscular  system  given  in  anatomical  text  books." 
A  single  body  presented  the  extraordinary  number  of  twenty-five 
distinct  abnormalities.  The  same  muscle  sometimes  varies  in 
many  ways:  thus  Prof.  Macalister  describes''  no  less  than  twenty 
distinct  variations  in  the  palmaris  accessorius. 

The  famous  old  anatomist,  Wolff,^  insists  that  the  internal 
viscera  are  more  variable  than  the  external  parts:  Nulla  parti- 
cula  est  quse  non  aliter  et  aliter  in  aliis  se  habeat  hominibus.  He 
has  even  written  a  treatise  on  the  choice  of  typical  examples  of 
the  viscera  for  representation.  A  discussion  on  the  beau-ideal 
of  the  liver,  lungs,  kidneys,  &c.,  as  of  the  human  face  divine, 
sounds  strange  in  our  earsi 

The  variability  or  diversity  of  the  mental  faculties  in  men  of 
the  same  race,  not  to  mention  the  greater  differences  between 
the  men  of  distinct  races,  is  so  notorious  that  not  a  word  need 
here  be  said.  So  it  is  with  the  lower  animal's.  All  who  have  had 
charge  of  menageries  admit  this  fact,  and  we  see  it  plainly  in  our 
dogs  and  other  domestic  animals.  Brehm  especially  insists  that 
each  individual  monkey  of  those  which  he  kept  tame  ip  Africa 
had  its  own  peculiar  disposition  and  temper:  he  mentions  one 
baboon  remarkable  for  its  high  intelligence;  and  the  keepers  in 
the  Zoological  Gardens  pointed  out  to  me  a  monkey,  belonging 
to  the  New  World  division,  equally  remarkable  for  intelligence. 
Rengger,  also,  insists  on  the  diversity  in  the  various  mental  char- 
acters of  the  monkeys  of  the  same  species  which  he  kept  in  Para- 
guay; and  this  diversity,  as  he  adds,  is  partly  innate,  and  partly 
the  result  of  the  manner  in  which  they  have  been  treated  or 
educated.^ 


8  'Anatomy  of  the  Arteries,'  by  R.  Quain.    Preface,  vol.  i.  1844. 

*  'Transact.  Royal  Soc.  Edinburgrh,'  vol.  xxlv.  pp.  175,  189. 

"  'Proc.  Royal  Soc'  1867,  p.  544;  also  1868,  pp.  483,  524.  There  is  a  pre- 
vious paper,  1866,  p.  229. 

«  'Proc.  R.  Irish  Academy,'  vol.  x.  1868,  p.  141. 

'  'Act.  Acad.  St.  Petersburg-,'  1778,  part  ii.  p.  217. 

8  Brehm,  'Thierleben,'  B.  i.  s.  58,  87.  Reng-ger.  'Saugethiere  von  Para- 
guay,' s.  57, 


MANNER   OF  DEVELOPMENT.  27 

I  have  elsewhere^  so  fully  discussed  the  subject  of  Inheritance, 
that  I  need  here  add  hardly  anything.  A  greater  number  of  facts 
have  been  collected  with  respect  to  the  transmission  of  the  most 
trifling,  as  well  as  of  the  most  important  characters  in  man,  than 
in  any  of  the  lower  animals;  though  the  facts  are  copious  enough 
with  respect  to  the  latter.  So  in  regard  to  mental  qualities,  their 
transmission  is  manifest  in  our  dogs,  horses,  and  other  domestic 
animals.  Besides  special  tastes  and  habits,  general  intelligence, 
courage,  bad  and  good  temper,  &c.,  are  certainly  transmitted.  With 
man  we  see  similar  facts  in  almost  every  family;  and  we  now 
know,  through  the  admirable  labors  of  Mr.  Galton,^"  that  genius 
which  implies  a  wonderfully  complex  combination  of  high  facul- 
ties, tends  to  be  inherited;  and,  on  the  other  hand,  it  is  too  certain 
that  insanity  and  deteriorated  mental  powers  likewise  run  in 
families. 

With  respect  to  the  causes  of  variability,  we  are  in  all  cases 
very  ignorant;  but  we  can  see  that  in  man  as  in  the  lower  ani- 
mals, they  stand  in  some  relation  to  the  conditions  to  which  each 
species  has  been  exposed,  during  several  generations.  Domes- 
ticated animals  vary  more  than  those  in  a  state  of  nature;  and 
this  is  apparently  due  to  the  diversified  and  changing  nature  of 
the  conditions  to  which  they  have  been  subjected.  In  this  respect 
the  different  races  of  man  resemble  domesticated  animals,  and  so 
do  the  individuals  of  the  same  race,  when  inhabiting  a  very  wide 
area,  like  that  of  America.  We  see  the  influence  of  diversified 
conditions  in  the  more  civilized  nations;  for  the  members  belong- 
ing to  different  grades  of  rank,  and  following  different  occupa- 
tions, present  a  greater  range  of  character  than  do  the  members 
of  barbarous  nations.  But  the  uniformity  of  savages  has  often 
been  exaggerated,  and  in  some  cases  can  hardly  be  said  to  exist." 
It  is,  nevertheless,  an  error  to  speak  of  man,  even  if  we  look  only 
to  the  conditions  to  which  he  has  been  exposed,  as  "far  more 
domesticated"^^  than  any  other  animal.  Some  savage  races,  such 
as  the  Australians,  are  not  exposed  to  more  diversified  conditions 
than  are  many  species  which  have  a  wide  range.  In  another  and 
much   more   important   respect,    man    differs    widely   from   any 

» 'Variation  of  Animals  and  Plants  under  pomestication,'  vol.  ii. 
chap.  xii. 

18  'Hereditary  Genius:  an  Inquiry  into  its  Laws  and  Consequences,' 
1869. 

^  Mr.  Bates  remarks  ('The  Naturalist  on  the  Amazons,'  1863,  vol.  ii. 
p.  159),  with  respect  to  the  Indians  of  the  same  South  American  tribe, 
"no  two  of  them  were  at  all  similar  in  the  shape  of  the  head;  one  man 
had  an  oval  visage  with  fine  features,  and  another  was  quite  Mongol- 
ian in  breadth  and  prominence  of  cheek,  spread  of  nostrils,  and 
obliquity  of  eyes." 

*2  Blumenbach,  'Treatises  on  Anthropolog.'  Eng.  translat.,  1865,  p.  205, 


2g  THE  DESCENT  OF  MAN. 

Strictly  domesticated  animal;  for  his  breeding  has  never  long 
been  controlled,  either  by  methodical  or  unconscious  selection. 
No  race  or  body  of  men  has  been  so  completely  subjugated  by 
other  men,  as  that  certain  individuals  should  be  preserved,  and 
thus  unconsciously  selected,  from  somehow  excelling  in  utility 
to  their  masters.  Nor  have  certain  m.ale  and  female  individuals 
been  intentionally  picked  out  and  matched,  except  in  the  well- 
known  case  of  the  Prussian  grenadiers;  and  in  this  case  man 
obeyed,  as  might  have  been  expected,  the  lav/  of  methodical  selec- 
tion; for  it  is  asserted  that  many  tall  men  were  reared  in  the 
villages  inhabited  by  the  grenadiers  and  their  tall  wives.  In 
Sparta,  also,  a  form  of  selection  was  follov/ed,  for  it  was  enacted 
that  all  children  should  be  examined  shortly  after  birth;  the  well- 
formed  and  vigorous  being  preserved,  the  others  left  to  perish.^^* 

If  we  consider  all  the  races  of  man  as  forming  a  single  species, 

I  his  range  is  enormous;  but  some  separate  races,  as  the  Americans 
?  and  Polynesians,  have  very  wide  ranges.  It  is  a  well-known 
I  law  that  widely-ranging  species  are  much  more  variable  than 
\  species  with  restricted  ranges;  and  the  variability  of  man  may 
with  more  truth  be  compared  v/ith  that  of  widely-ranging  species, 
than  with  that  of  domesticated  animals. 

Not  only  does  variability  appear  to  be  induced  in  man  and 
the  lower  animals  by  the  same  general  causes,  but  in  both  the 

13  Mitford's  'History  of  Greece,'  vol.  i.  p.  282.  It  appears  also  from 
a  passage  in  Xenophon's  'Memorabilia,'  B.  ii.  4  (to  which  my  attention 
has  been  called  by  the  Rev.  J.  N.  Hcare),  that  it  was  a  well  recognized 
principle  with  the  Greeks,  that  men  ought  to  select  their  wives  with  a 
view  to  the  health  and  vigor  of  their  children.  The  Grecian  poet, 
Theognis,  who  lived  550  B.  C,  clearly  saw  how  important  selection,  if 
carefully  applied,  would  be  for  the  improvement  of  mankind.  He  saw, 
likewise,  that  wealth  often  checks  the  proper  action  of  sexual  selec- 
tion.   He  thus  writes: 

"With  kine  and  horses,  Kurnus!    we  proceed 
By  reasonable  rules,  and  choose  a  breed 
For  profit  and  increase,  at  any  price; 
Of  a  sound  stock,  without  defect  or  vice. 
But,  in  the  daily  matches  that  we  make, 
The  price  is  everything:    for  money's  sake. 
Men  marry:    women  are  in  marriage  given; 
The  churl  or  ruffian,  that  in  wealth  has  thriven. 
May  match  his  offspring  with  the  proudest  race: 
Thus  everything  is  mix'd,  noble  and  base! 
If  then  in  outward  manner,  form,  and  mind. 
You  find  us  a  degraded  motley  kind. 
Wonder  no  more,  my  friend!  the  cause  is  plain, 
And  to  lament  the  consequence  is  vain." 
(The  Works  of  J.  Hookham  Frere,  vol.  ii.  1872,  p.  334.) 


MANNER   OF  DEVELOPMENT.  29 

same  parts  of  the  body  are  affected  in  a  closely  analogous  man- 
ner. This,  has  been  proved  in  such  full  detail  by  Godron  and 
Quatrefages,  that  I  need  here  only  refer  to  their  works."  Mon- 
strosities, which  graduate  Into  slight  variations,  are  likewise  so 
similar  in  man  and  the  lower  animals,  that  the  same  classification 
and  the  same  terms  can  be  used  for  both,  as  has  been  shown  by 
Isidore  Geoffroy  St.-Hilaire.^^  In  my  work  on  the  variation  of 
domestic  animals,  I  have  attempted  to  arrange  in  a  rude  fashion 
the  laws  of  variation  under  the  following  heads:— The  direct  and 
definite  action  of  changed  conditions,  as  exhibited  by  all  or  nearly 
all  the  individuals  of  the  same  species,  varying  in  the  same  man- 
ner under  the  same  circumstances.  The  effects  of  the  long-con- 
tinued use  or  disuse  of  parts.  The  cohesion  of  homologous  parts. 
The  variability  of  multiple  parts.  Compensation  of  growth;  but 
of  this  law  I  have  found  no  good  instance  in  the  case  of  man.  The 
effects  of  the  mechanical  pressure  of  one  part  on  another;  as  of 
the  pelvis  on  the  cranium  of  the  infant  in  the  womb.  Arrests  of 
development,  leading  to  the  diminution  or  suppression  of  parts. 
The  reappearance  of  long-lost  characters  through  reversion.  And 
lastly,  correlated  variation.  All  these  so-called  laws  apply  equally 
to  man  and  the  lower  animals;  and  most  of  them  even  to  plants. 
It  would  be  superfluous  here  to  discuss  all  of  them;"  but  several 
are  so  important,  that  they  must  be  treated  at  considerable  length. 

The  direct  and  definite  action  of  changed  conditions. — This  is  a 
most  perplexing  subject.  It  cannot  be  denied  that  changed  con- 
ditions produce  some,  and  occasionally  a  considerable  effect,  on 
organisms  of  all  kinds;  and  it  seems  at  first  probable  that  if  suf- 
ficient time  were  allowed  this  would  be  the  invariable  result. 
But  I  have  failed  to  obtain  clear  evidence  in  favor  of  this  con- 
clusion; and  valid  reasons  may  be  urged  on  the  other  side,  at 
least  as  far. as  the  innumerable  structures  are  concerned,  which 
are  adapted  for  special  ends.  There  can,  however,  be  no  doubt 
that  changed  conditions  induce  an  almost  indefinite  amount  of 
fluctuating  variability,  by  which  the  whole  organization  is  ren- 
dered in  some  degree  plastic. 

In  the  United  States,  above  1,000,000  soldiers,  who  served  in 

1*  Godron,  'De  I'Espece,'  1859,  torn.  ii.  livre  3.  Quatrefages,  'Unite  de 
I'Espece  Humaine,'  1881.  Also  Lectures  on  Anthropology,  given  in  the 
'Revue  des  Cours  Scientifiques'  1866-1868. 

15  'Hist.  Gen.  et  Part,  des  Anomalies  de  1' Organisation'  in  three  vol- 
umes, torn.  i.  1832. 

i«  I  have  fully  discussed  these  laws  in  my  'Variation  of  Animals  and 
Plants  under  Domestication,'  vol.  ii.  chap.  xxii.  and  xxiii.  M.  J.  P. 
Durand  has  lately  (1868)  published  a  valuable  essay  'De  I'lnfluence  des 
Milieux,'  &c.  He  lays  much  stress,  in  the  case  of  plants,  on  the  nature 
of  the  soil. 


3d  THE  DESCENT  OF  MAN. 

the  late  war,  were  measured,  and  the  States  in  which  they  were 
born  and  reared  were  recorded."  From  this  astonishing  number 
of  observations  it  is  proved  that  local  influences  of  some  kind 
act  directly  on  stature;  and  we  further  learn  that  "the  State 
"where  the  physical  growth  has  in  great  measure  taken  place, 
"and  the  State  of  birth,  which  indicates  the  ancestry,  seem  to 
"exert  a  marked  influence  on  the  stature."  For  instance,  it  is 
established,  "that  residence  in  the  'Western  States,  during  the 
"years  of  growth,  tends  to  produce  increase  of  stature."  On  the 
other  hand,  it  is  certain  that  with  sailors,  their  life  delays  growth, 
as  shown  "by  the  great  difference  between  the  statures  of  soldiers 
"and  sailors  at  the  ages  of  seventeen  and  eighteen  years."  Mr.  B. 
A.  Gould  endeavored  to  ascertain  the  nature  of  the  influences 
which  thus  act  on  stature;  but  he  arrived  only  at  negative  results, 
namely,  that  they  did  not  relate  to  climate,  the  elevation  of  the 
land,  soil,  nor  even  "in  any  controlling  degree"  to  the  abundance 
or  the  need  of  the  comforts  of  life.  This  latter  conclusion  is 
directly  opposed  to  that  arrived  at  by  Villerme,  from  the  statistics 
of  the  height  of  the  conscripts  in  different  parts  of  France.  When 
we  compare  the  differences  in  stature  between  the  Polynesian 
chiefs  and  the  lower  orders  within  the  same  islands,  or  between 
the  inhabitants  of  the  fertile  volcanic  and  low  barren  coral  islands 
of  the  same  ocean,^^  or  again  between  the  Fuegians  on  the  eastern 
and  western  shores  of  their  country,  where  the  means  of  subsist- 
ence are  very  different,  it  is  scarcely  possible  to  avoid  the  con- 
clusion that  better  food  and  greater  comfort  do  influence  stature. 
But  the  preceding  statements  show  how  difiicult  it  is  to  arrive 
at  any  precise  result.  Dr.  Beddoe  has  lately  proved  that,  with 
the  inhabitants  of  Britain,  residence  in  towns  and  certain  occupa- 
tions have  a  deteriorating  influence  on  height;  and  he  infers  that 
the  result  is  to  a  certain  extent  inherited,  as  is  likewise  the  case 
in  the  United  States.  Dr.  Beddoe  further  believes  that  wherever 
a  "race  attains  its  maximum  of  physical  development,  it  rises 
"highest  in  energy  and  moral  vigor."^" 

Whether  external  conditions  produce  any  other  direct  effect 
on  man  is  not  known.  It  might  have  been  expected  that  dif- 
ferences of  climate  would  have  had  a  marked  influence,  in  as  much 
as  the  lungs  and  kidneys  are  brought  into  activity  under  a  low 

"  'Investigations  in  Military  and  Anthrop.  Statistics,'  &c.  1869,  by  B. 
A.  Gould,  p.  93,  107,  126,  131,  134. 

18  For  the  Polynesians,  see  Prichard's  'Physical  Hist,  of  Mankind,' 
vol.  V.  1847,  p.  145,  283.  Also  Godron,  'De  I'Espece,'  torn.  ii.  p.  289.  There 
is  also  a  remarkable  difference  in  appearance  between  the  closely-allied 
Hindoos  inhabiting  the  Upper  Ganges  and  Bengal;  see  Elphinstone's 
'History  of  India,'  vol.  i.  p.  324. 

19  'Memoirs,  Anthropolog.  Soc.  vol.  ill.  1867-1S69.  pp.  561,  565,  567. 


MANNER   OF  DE;VEL0PMENT.  31 

temperature,  and  the  liver  and  skin  under  a  high  one.^"  It  was 
formerly  thought  that  the  color  of  the  skin  and  the  character 
of  the  hair  were  determined  hy  light  or  heat;  and  although  it 
can  hardly  be  denied  that  some  effect  is  thus  produced,  almost 
all  observers  now  agree  that  the  effect  has  been  very  small,  even 
after  exposure  during  many  ages.  But  this  subject  will  be  more 
properly  discussed  when  we  treat  of  the  different  races  of  man- 
kind. With  our  domestic  animals  there  are  grounds  for  believing 
that  cold  and  damp  directly  affect  the  growth  of  the  hair;  but 
I  have  not  met  with  any  evidence  on  this  head  in  the  case  of 
man. 

Effects  of  the  increased  Use  and  Disuse  of  Parts. — It  is  well 
known  that  use  strengthens  the  muscles  in  the  individual,  and 
complete  disuse,  or  the  destruction  of  the  proper  nerve,  weakens 
them.  When  the  eye  is  destroyed,  the  optic  nerve  often  becomes 
atrophied.  When  an  artery  is  tied,  the  lateral  channels  increase 
not  only  in  diameter,  but  in  the  thickness  and  strength  of  their 
coats.  When  one  kidney  ceases  to  act  from  disease,  the  other 
increases  in  size,  and  does  double  work.  Bones  increase  not 
only  in  thickness,  but  in  length,  from  carrying  a  greater  weight.^^ 
Different  occupations,  habitually  followed,  lead  to  changed  pro- 
portions in  various  parts  of  the  body.  Thus  it  was  ascertained 
by  the  United  States  Commission^-  that  the  legs  of  the  sailors 
employed  in  the  late  war  were  longer  by  0.217  of  an  inch  than 
those  of  the  soldiers,  though  the  sailors  were  on  an  average 
shorter  men;  whilst  their  arms  were  shorter  by  1.09  of  an  inch, 
and  therefore,  out  of  proportion,  shorter  in  relation  to  their  lesser 
height.  This  shortness  of  the  arms  is  apparently  due  to  their 
greater  use,  and  is  an  unexpected  result:  but  sailors  chiefly  use 
their  arms  in  pulling,  and  not  in  supporting  weights.  With 
sailors,  the  girth  of  the  neck  and  the  depth  of  the  instep  are 
greater,  whilst  the  circumference  of  the  chest,  waist,  and  hips  is 
less,  than  in  soldiers. 

Whether  the  several  foregoing  modifications  would  become 
hereditary,  if  the  same  habits  of  life  were  followed  during  many 
generations,  is  not  known,  but  it  is  probable.  Rengger-^  attributes 
the  thin  legs  and  thick  arms  of  the  Payaguas  Indians  to  successive 
generations  having  passed  nearly  their  whole  lives  in  canoes, 

20  Dr.  Brakenridg-e,  'Theory  of  Diathesis,'  'Medical  Times,'  June  19 
and  July  17,  1869. 

21 1  have  given  authorities  for  these  several  statements  in  my  'Varia- 
tion of  Animals  under  Domestication,'  vol.  ii.  pp.  297-300.  Dr.  Jaeger, 
"Ueber  das  Langenwachsthum  der  Knbchen,"  'Jenaischen  Zeitschrift,' 
B.  V.  Heft  i. 

22  'Investigations,'  &c.    By  B.  A.  Gould,  1869,  p.  288. 

23  'Saugethiere  von  Paraguay,'  1830,  s.  4. 


32  THE  DESCENT  OF  MAN. 

with  their  lower  extremities  motionless.  Other  writers  have 
come  to  a  similar  conclusion  in  analogous  cases.  According  to 
Cranz,-*  who  lived  for  a  long  time  with  the  Esquimaux,  "the  na- 
"tives  believe  that  ingenuity  and  dexterity  in  seal-catching  (their 
"highest  art  and  virtue)  is  hereditary;  there  is  really  something 
*'in  it,  for  the  son  of  a  celebrated  seal-catcher  will  distinguish 
"himself,  though  he  lost  his  father  in  childhood."  But  in  this 
case  it  is  mental  aptitude,  quite  as  much  as  bodily  structure,  which 
appears  to  be  inherited.  It  is  asserted  that  the  hands  of  English 
laborers  are  at  birth  larger  than  those  of  the  gentry."^  From  the 
correlation  which  exists,  at  least  in  some  cases,^°  between  the  de- 
velopment of  the  extremities  and  of  the  jaws,  it  is  possible  that 
in  those  classes  which  do  not  labor  much  with  their  hands  and 
feet,  the  jaws  would  be  reduced  in  size  from  this  cause.  That  they 
are  generally  smaller  in  refined  and  civilized  men  than  in  hard- 
working men  or  savages,  is  certain.  But  with  savages,  as  Mr. 
Herbert  Spencer^^  has  remarked,  the  greater  use  of  the  jaws  in 
chewing  coarse,  uncooked  food,  would  act  in  a  direct  manner  on 
the  masticatory  muscles,  and  on  the  bones  to  which  they  are  at- 
tached. In  infants,  long  before  birth,  the  skin  on  the  soles  of 
the  feet  is  thicker  than  on  any  other  part  of  the  body;^^  and  it 
can  hardly  be  doubted  that  this  is  due  to  the  inherited  effects  of 
pressure  during  a  long  series  of  generations. 

It  is  familiar  to  every  one  that  watchmakers  and  engravers 
are  liable  to  be  short-sighted,  whilst  men  living  much  out  of 
doors,  and  especially  savages,  are  generally  long-sighted.^^  Short- 
sight  and  long-sight  certainly  tend  to  be  inherited.^"  The  infe- 
riority of  Europeans,  in  comparison  with  savages,  in  eyesight  and 
in  the  other  senses,  is  no  doubt  the  accumulated  and  transmitted 
effect  of  lessened  use  during  many  generations;    for  Rengger^^ 

2*  'History  of  Greenland,'  Eng-.  translat.  1767,  vol.  i.  p.  280. 

25  'Intermarriage.'    By  Alex.  Walker,  1838,  p.  377. 

26  'The  Variation  of  Animals  under  Domestication,'  vol.  i.  p.  173. 

27  'Principles  of  Biology,'  vol.  i.  p.  455. 

28  Paget,  'Lectures  on  Surgical  Pathology,'  vol.  ii.  1853,  p.  209. 

29  It  is  a  singular  and  unexpected  fact  that  sailors  are  inferior  to 
landsmen  in  their  mean  distance  of  distinct  vision.  Dr.  B.  A.  Gould 
('Sanitary  Memoirs  of  the  War  of  the  Rebellion,'  1869,  p.  530),  has  proved 
this  to  be  the  case;  and  he  accounts  for  it  by  the  ordinary  range  of 
vision  in  sailors  being  "restricted  to  the  length  of  the  vessel  and  the 
height  of  the  masts." 

30  'The  variation  of  Animals  Under  Domestication,'  vol.  i.  p.  8. 

31  'Saugethiere  von  Paraguay,'  s.  8,  10.  I  have  had  good  opportuni- 
ties for  observing  the  extraordinary  power  of  eyesight  in  the  Fuegians. 
See  also  Lawrence  ('Lectures  on  Physiology,'  &c.,  1822,  p.  404)  on  this 
same  subject.  M.  Giraud-Teulon  has  recently  collected  ('Revue  des 
Cours  Scientifiques,'  1870,  p.  625)  a  large  and  valuable  body  of  evidence 
proving  that  the  cause  of  short-sight,  "C'est  le  travail  assidu.  de  pres," 


MANNER    OF   DEVELOPMENT.  33 

states  that  he  has  repeatedly  observed  Europeans,  who  had  been 
brought  up  and  spent  their  whole  lives  with  the  wild  Indians, 
who  nevertheless  did  not  equal  them  in  the  sharpness  of  their 
senses.  The  same  naturalist  observes  that  the  cavities  in  the 
Skull  for  the  reception  of  the  several  sense-organs  are  larger  in 
the  American  aborigines  than  in  Europeans;  and  this  probably 
indicates  a  corresponding  difference  in  the  dimensions  of  the  or- 
gans themselves.  Blumenbach  has  also  remarked  on  the  large 
size  of  the  nasal  cavities  in  the  skulls  of  the  American  aborigines, 
and  connects  this  fact  with  their  remarkably  acute  power  of  smell. 
The  Mongolians  of  the  plains  of  Northern  Asia,  according  to 
Pallas,  have  wonderfully  perfect  senses;  and  Prichard  believes 
that  the  great  breadth  of  their  skulls  across  the  zygomas  follows 
from  their  highly-developed  sense-organs.^- 

The  Quechua  Indians  inhabit  the  lofty  plateaux  of  Peru;  and 
Alcide  d'Orbigny  states'-  that  from  continually  breathing  a  highly 
rarefied  atmosphere,  they  have  acquired  chests  and  lungs  of  ex- 
traordinary dimensions.  The  cells,  also,  of  the  lungs  are  larger 
and  more  numerous  than  in  Europeans.  These  observations  have 
been  doubted;  but  Mr.  D.  Forbes  carefully  measured  many  Ay- 
maras,  an  allied  race,  living  at  the  height  of  between  10,000  and 
15,000  feet;  and  he  informs  me^^  that  they  differ  conspicuously 
from  the  men  of  all  other  races  seen  by  him  in  the  circumference 
and  length  of  their  bodies.  In  his  table  of  measurements,  the 
stature  of  each  man  is  taken  at  1000,  and  the  other  measurements 
are  reduced  to  this  standard.  It  is  here  seen  that  the  extended 
arms  of  the  Aymaras  are  shorter  than  those  of  Europeans,  and 
much  shorter  than  these  of  Negroes.  The  legs  are  likewise  shorter; 
and  they  present  this  remarkable  peculiarity,  that  in  every  Ay- 
mara  measured,  the  femur  is  actually  shorter  than  the  tibia.  On 
an  average,  the  length  of  the  femur  to  that  of  the  tibia  is  as  211 
to  252;  whilst  in  two  Europeans,  measured  at  the  same  time,  the 
femora  to  the  tibise  were  at  244  to  230;  and  in  three  Negroes  as 
258  to  241.  The  humerus  is  likewise  shorter  relatively  to  the  fore- 
arm. This  shortening  of  that  part  of  the  limb  which  is  nearest  to 
the  body,  appears  to  be,  as  suggested  to  me  by  Mr.  Forbes,  a  case 
of  compensation  in  relation  with  the  greatly  increased  length  of 
the  trunk.  The  Aymaras  present  some  other  singular  points  of 
structure,  for  instance,  the  very  small  projection  of  the  heel. 

These  men  are  so  thoroughly  acclimatized  to  their  cold  and 
lofty  abode,  that  when  formerly  carried  down  by  the  Spaniards 


32  Prichard,   'Phys.  Hist,  of  Mankind,'  on  the  authority  of  Blumen- 
bach, vol.  i.  1851,  p.  311;    for  the  statement  by  Pallas,  vol.  iv.  1844,  p.  407. 

33  Quoted  by  Prichard,  'Researches  into  the  Phys.  Hist,  of  Mankind,' 
vol.  V.  p.  463. 

34  Mr.   Forbes'   valuable  paper  is  now  published  in  the    'Journal  of 
the  Ethnological  Soc.  of  London,'  new  series,  vol.  ii.  1870,  p.  193. 

4 


34  THE  DESCENT  OF  MAN. 

to  the  low  eastern  plains,  and  when  now  tempted  down  by  high 
wages  to  the  gold-washings,  they  suffer  a  frightful  rate  of  mor- 
tality. Nevertheless  Mr.  Forbes  found  a  few  pure  families  which 
had  survived  during  two  generations:  and  he  observed  that  they 
still  inherited  their  characteristic  peculiarities.  But  it  was  mani- 
fest, even  without  measurement,  that  these  peculiarities  had  all 
decreased;  and  on  measurement,  their  bodies  were  found  not  to  be 
so  much  elongated  as  those  of  the  men  on  the  high  plateau;  whilst 
their  femora  had  become  somewhat  lengthened,  as  had  their  tibiae, 
although  in  a  less  degree.  The  actual  measurements  may  be  seen 
by  consulting  Mr.  Forbes's  memoir.  From  these  observations, 
there  can,  I  think,  be  no  doubt  that  residence  during  many  genera- 
tions at  a  great  elevation  tends,  both  directly  and  indirectly,  to 
induce  inherited  modifications  in  the  proportions  of  the  body." 

Although  man  may  not  have  been  much  modified  during  the 
latter  stages  of  his  existence  through  the  increased  or  decreased 
use  of  parts,  the  facts  now  given  show  that  his  liability  in  this 
respect  has  not  been  lost;  and  we  positively  know  that  the  same 
law  holds  good  with  the  lower  animals.  Consequently  we  may 
infer  that  when  at  a  remote  epoch  the  progenitors  of  man  were 
in  a  transitional  state,  and  were  changing  from  quadrupeds  into 
bipeds,  natural  selection  would  probably  have  been  greatly  aided 
by  the  inherited  effects  of  the  increased  or  diminished  use  of  the 
different  parts  of  the  body. 

Arrests  of  Development.  — There  is  a  difference  between  arrested 
development  and  arrested  growth,  for  parts  in  the  former  state 
continue  to  grow  whilst  still  retaining  their  early  condition. 
Various  monstrosities  come  under  this  head;  and  some,  as  a 
cleft-palate,  are  known  to  be  occasionally  inherited.  It  will 
suffice  for  our  purpose  to  refer  to  the  arrested  brain-development 
of  microcephalous  idiots,  as  described  in  Vogt's  memoir.^^  Their 
skulls  are  smaller,  and  the  convolutions  of  the  brain  are  less  com- 
plex than  in  normal  men.  The  frontal  sinus,  or  the  projection 
over  the  eyebrows,  is  largely  developed,  and  the  jaws  are  prog- 
nathous to  an  "effrayant"  degree,  so  that  these  idiots  somewhat 
resemble  the  lower  types  of  mankind.  Their  intelligence,  and 
most  of  their  mental  faculties,  are  extremely  feeble.  They 
cannot  acquire  the  power  of  speech,  and  are  wholly  incapable  of 
prolonged  attention,  but  are  much  given  to  imitation.  They  are 
strong  and  remarkably  active,  continually  gamboling  and  jumping 
about,  and  making  grimaces.  They  often  ascend  stairs  on  all- 
fours;     and  are  curiously  fond  of  climbing  up  furniture  or  trees. 

35  Dr.  Wilckens  ('Landwirthschaft.  Wochenblatt,'  No.  10,  1869)  has 
lately  published  an  interesting  Essay  showing-  how  domestic  animals, 
which  live  in  mountainous  regions,  have  their  frames  modified. 

36  'Memoire  sur  les  Microcephales,'  1867,  pp.  50,  125,  169,  171,  184-198. 


MANNER   OF  DEVELOPMENT.  35 

We  are  thus  reminded  of  the  delight  shown  by  almost  all  boys 
in  climbing  trees;  and  this  again  reminds  us  how  lambs  and  kids, 
originally  alpine  animals,  delight  to  frisk  on  any  hillock,  however 
small.  Idiots  also  resemble  the  lower  animals  in  some  other 
respects;  thus  several  cases  are  recorded  of  their  carefully  smell- 
ing every  mouthful  of  food  before  eating  it.  One  idiot  is  described 
as  often  using  his  mouth  in  aid  of  his  hands,  whilst  hunting  for 
lice.  They  are  often  filthy  in  their  habits,  and  have  no  sense  of 
decency;  and  several  cases  have  been  published  of  their  bodies 
being  remarkably  hairy.'^ 

Reversion. —Many  of  the  cases  to  be  here  given,  might  have 
been  introduced  under  the  last  heading.  When  a  structure  is 
arrested  in  its  development,  but  still  continues  growing,  until  it 
closely  resembles  a  corresponding  structure  in  some  lower  and 
adult  member  of  the  same  group,  it  may  in  one  sense  be  con- 
sidered as  a  case  of  reversion.  The  lower  members  in  a  group 
give  us  some  idea  how  the  common  progenitor  was  probably  con- 
structed; and  it  is  hardly  credible  that  a  complex  part,  arrested 
at  an  early  phase  of  embryonic  development,  should  go  on  grow- 
ing so  as  ultimately  to  perform  its  proper  function,  unless  it  had 
acquired  such  power  during  some  earlier  state  of  existence,  when 
the  present  exceptional  or  arrested  structure  was  normal.  The 
simple  brain  of  a  microcephalous  idiot,  in  as  far  as  it  resembles 
that  of  an  ape,  may  in  this  sense  be  said  to  offer  a  case  of  rever- 
sion.^^   There  are  other  cases  which  come  more  strictly  under  our 

87  Prof.  Laycock  sums  up  the  character  of  brute-like  idiots  by  calling- 
them  "theroid;"  'Journal  of  Mental  Science,'  July,  1863.  Dr.  Scott  ('The 
Deaf  and  Dumb,'  2nd  edit.,  1870,  p.  10)  has  often  observed  the  imbecile 
smelling-  their  food.  See,  on  Uiis  same  subject,  and  on  the  hairiness 
of  idiots.  Dr.  Maudsley,  'Body  and  Mind,'  1870,  pp.  46-51.  Pinei  has  also 
given  a  striking  case  of  hairiness  in  an  idiot. 

38  In  my  'Variation  of  Animals  under  Domestication'  (vol.  ii.  p.  57), 
I  attributed  the  not  very  rare  cases  of  supernumerary  mammae  in 
women  to  reversion.  I  was  led  to  this  as  a  probable  conclusion,  by  the 
additional  mammae  being-  g-enerally  placed  symmetrically  on  the 
breast;  and  more  especially  from  one  case,  in  which  a  single  efficient 
mamma  occurred  in  the  inguinal  region  of  a  woman,  the  daughter  of 
another  woman  with  supernumerary  mammae.  But  I  now  find  (see, 
for  instance.  Prof.  Preyer,  'Der  Kampf  um  das  Dasein,'  1859,  s.  45)  that 
mammae  erraticae  occur  in  other  situations,  as  on  the  back,  in  the 
armpit,  and  on  the  thigh;  the  mammae  in  this  latter  instance  having 
given  so  much  milk  that  the  child  was  thus  nourished.  The  probability 
that  the  additional  mammae  are  due  to  reversion  is  thus  much  weak- 
ened; nevertheless  it  still  seems  to  me  probable,  because  tv/o  pairs 
are  often  found  symmetrically  on  the  breast;  and  of  this  I  myself 
have  received  information  in  several  cases.  It  is  well  known  that  some 
Lemurs  normally  have  two  pairs  of  mammae  on  the  breast.  Five 
cases  have  been  recorded  of  the  presence  of  more  than  a  pair  of  mam- 


3^  THE  DESCENT  OF  MAN. 

present  head  of  reversion.  Certain  structures,  regularly  occurring 
in  tlie  lower  members  of  the  group  to  which  man  belongs,  occa- 
sionally make  their  appearance  in  him,  though  not  found  in  the 
normal  human  embryo;  or,  if  normally  present  in  the  human 
embryo,  they  become  abnormally  developed,  although  in  a  manner 
which  is  normal  in  the  lower  members  of  the  group.  These  re- 
marks will  be  rendered  clearer  by  the  following  illustrations. 

In  various  mammals  the  uterus  graduates  from  a  double  organ, 
with  two  distinct  orifices  and  two  passages,  as  in  the  marsupials, 
into  a  single  organ,  which  is  in  no  way  double,  except  from 
having  a  slight  internal  fold,  as  in  the  higher  apes  and  man. 

mae  (of  course  rudimentary)  in  the  male  sex  of  mankind;  see  'Journal 
of  Anat.  and  Physiology,'  1872,  p.  56,  for  a  case  given  by  Dr.  Handj^side, 
in  which  two  brothers  exhibited  this  peculiarity;  see  also  a  paper  by 
Dr.  Bartels  in  'Reichert's  and  du  Bois-Reymond's  Archiv.,'  1872,  p.  304. 
In  one  of  the  cases  alluded  to  by  Dr.  Bartels,  a  man  bore  five  mam- 
mae, one  being-  medial  and  placed  above  the  navel;  Meckel  von  Hems- 
bach  thinks  that  this  latter  case  is  illustrated  by  a  medial  mamma 
occurring  in  certain  Cheiroptera.  On  the  whole  we  may  well  doubt  if 
additional  mammae  v/ould  ever  have  been  developed  in  both  sexes  of 
mankind,  had  not  his  early  progenitors  been  provided  with  more  than 
a  single  pair. 

In  the  above  work  (vol.  ii.  p.  12),  I  alSo  attributed,  though  with  much 
hesitation,  the  frequent  cases  of  polydactylism  in  men  and  various  ani- 
mals to  reversion.    I  was  partly  led  to  this  through  Prof.  Owen's  state- 
ment, that  some  of  the  Ichthyopterygia  possess  more  than  five  digits, 
and  therefore,  as  I  supposed,  had  retained  a  primordial  condition;    but 
Prof.   Gegenbaur   ('Jenaischen  Zeitschrift,'   B.   v.   Heft  3,  s.   341),   dis- 
putes Owen's  conclusion.    On  the  other  hand,  according  to  the  opinion 
lately  advanced  by  Dr.  Gunther,  on  the  paddle  of  Ceratodus,  which  is 
provided  with  articulated  bony  rays  on  both  sides  of  a  central  chain  of 
bones,  there  seems  no  great  difficulty  in  admitting  that  six  or  more 
digits  on  one  side,  or  on  both  sides,  might  reappear  through  reversion. 
I  am  informed  by  Dr.  Zouteveen  that  there  is  a  case  on  record  of  a 
man  having  twenty-four  fingers  and  twenty-four  toes!    I  v/as  chiefly 
led  to  the  conclusion  that  the  presence  of  supernumerary  digits  might 
be  due  to  reversion  from  the  fact  that  such  digits,  not  only  are  strongly 
inherited  but,   as  I  then  believed,   had  the  power  of  regrowth   a^ter 
amputation,  like  the  norm.al  digits  of  the  lower  vertebrata.    But  I  have 
explained  in  the  Second  Edition  of  my  Variation  under  Domestication 
why  I  now  place  little  reliance  on  the  recorded  cases  of  such  regrowth. 
Nevertheless  it  deserves  notice,  in  as  much  as  arrested  developm^ent 
and  reversion  are  intimately  related  processes;   that  various  structures 
m  an  embryonic  or  arrested  condition,   such  as   a  cleft  palate,  bifid 
uterus,  &c.,  are  frequently  accompanied  by  polydactylism.     This  has 
been  strongly  insisted  on  by  Meckel  and  Isidore  Geoffrey  St.  Hilaire. 
But  at  present  it  is  the  safest  course  to  give  up  altogether  the  idea 
that  there  is  any  relation  between  the  development  of  supernumerary 
digits  and  reversion  to  some  lowly  organized  progenitor  of  man. 


MANNER    OF   DEVELOPMENT.  37 

The  rodents  exhibit  a  perfect  series  of  gradations  between  these 
two  extreme  states.  In  all  mammals  the  uterus  is  developed  frons 
two  simple  primitive  tubes,  the  inferior  portions  of  which  form 
the  cornua;  and  it  is,  in  the  words  of  Dr.  Parre,  "by  the  coa- 
"lescence  of  the  two  cornua  at  their  lower  extremities  that  the 
"body  of  the  uterus  is  formed  in  man;  while  in  those  animals 
"in  which  no  middle  portion  of  body  exists,  the  cornua  remain 
"un-united.  As  the  development  of  the  uterus  proceeds,  the  two 
"cornua  become  gradually  shorter,  until  at  length  they  are  losty 
"or,  as  it  were,  absorbed  into  the  body  of  the  uterus."  The  angler 
of  the  uterus  are  still  produced  into  cornua,  even  in  animals  aa 
high  up  in  the  scale  as  the  lower  apes  and  lemurs. 

Now  in  women,  anomalous  cases  are  not  very  infrequent,  in 
which  the  mature  uterus  is  furnished  with  cornua,  or  is  partially 
divided  into  two  organs;  and  such  cases,  according  to  Owen, 
repeat  "the  grade  of  concentrative  development,"  attained  by 
certain  rodents.  Here  perhaps  we  have  an  instance  of  a  simple 
arrest  of  embryonic  development,  with  subsequent  growth  and 
perfect  functional  development;  for  either  side  of  the  partially 
double  uterus  is  capable  of  performing  the  proper  office  of 
gestation.  In  other  and  rarer  cases,  two  distinct  uterine  cavities 
are  formed,  each  having  its  proper  orifice  and  passage.^  No 
such  stage  is  passed  through  during  the  ordinary  development 
of  the  embryo,  and  it  is  difficult  to  believe,  though  perhaps  not 
impossible,  that  the  two  simple,  minute,  primitive  tubes  should 
know  how  (if  such  an  expression  may  be  used)  to  grow  into  two 
distinct  uteri,  each  with  a  well-constructed  orifice  and  passage^ 
and  each  furnished  with  numerous  muscles,  nerves,  glands  and 
vessels,  if  they  had  not  formerly  passed  through  a  similar  course 
of  development,  as  in  the  case  of  existing  marsupials.  No  one 
will  pretend  that  so  perfect  a  structure  as  the  abnormal  double 
uterus  in  woman  could  be  the  result  of  mere  chance.  But  the 
principle  of  reversion,  by  which  a  long-lost  structure  is  called 
back  into  existence,  might  serve  as  the  guide  for  its  full  develop- 
ment, even  after  the  lapse  of  an  enormous  interval  of  time. 

Professor  Canestrini,  after  discussing  the  foregoing  and  various 
analogous  cases,  arrives  at  the  same  conclusion  as  that  just  given. 
He  adduces  another  instance,  in  the  case  of  the  malar  bone,*" 

3»  See  Dr.  A.  Farre's  well-known  article  in  the  'Cyclopaedia  of  Anat- 
omy and  Physiolog-y,'  vol.  v.,  1859,  p.  642.  Ov/en,  'Anatomy  of  Verte- 
brates,' vol.  iii.,  1868,  p.  687.  Professor  Turnier  in  'Edinburgh  Medical 
Journal,'  February,  1865. 

*° 'Annuario  della  Soc.  dei  Naturalist!  in  Modena,'  1867,  p.  83.  Prof. 
Canestrini  g-ives  extracts  on  this  subject  fro'm  various  authorities. 
Laurillard  remarks,  that  as  he  has  found  a  complete  similarity  in  the 
form,  proportions,  aaid  connection  of  the  two  malar  bones  in  several 
human  subjects  and  in  certain  apes,  he  cannot  consider  this  disposition 
4 


gg  THE  DESCENT  OF  MAN. 

which,  in  some  of  the  Quadrumana  and  other  mammals,  normally 
consists  of  two  portions.  This  is  its  condition  in  the  human  fcetus 
when  two  months  old;  and  through  arrested  development,  it 
sometimes  remains  thus  in  man  when  adult,  more  especially  in 
the  lower  prognathous  races.  Hence  Canestrini  concludes  that 
some  ancient  progenitor  of  man  must  have  had  this  bone  nor- 
mally divided  into  two  portions,  which  afterwards  became  fused 
together.  In  man  the  frontal  bone  consists  of  a  single  piece,  but 
in  the  embryo,  and  in  children,  and  in  almost  all  the  lower  mam- 
mals, it  consists  of  two  pieces  separated  by  a  distinct  suture.  This 
suture  occasionally  persists  more  or  less  distinctly  in  man  after 
maturity;  and  more  frequently  in  ancient  than  in  recent  crania, 
especially,  as  Canestrini  has  observed,  in  those  exhumed  from 
the  Drift,  and  belonging  to  the  brachycephalic  type.  Here  again 
he  comes  to  the  same  conclusion  as  in  the  analogous  case  of  the 
malar  bones.  In  this,  and  other  instances  presently  to  be  given, 
the  cause  of  ancient  races  approaching  the  lower  animals  in 
certain  characters  more  frequently  than  do  the  modern  races, 
appears  to  be,  that  the  latter  stand  at  a  somewhat  greater  distance 
in  the  long  line  of  descent  from  their  early  semi-human  progeni- 
tors. 

Various  other  anomalies  in  man,  more  or  less  analogous  to  the 
foregoing,  have  been  advanced  by  different  authors,  as  cases  of 
reversion;  but  these  seem  not  a  little  doubtful,  for  we  have  to 
descend  extremely  low  in  the  mammalian  series,  before  we  find 
such  structures  normally  present.*^ 


of  the  parts  as  simply  accidental.  Another  paper  on  this  same  anomaly 
has  been  published  by  Dr.  Saviotti  in  the  'Gazetta  delle  Cliniche'  Turin, 
1871,  where  he  says  that  traces  of  the  division  may  be  detected  in  about 
two  per  cent,  of  adult  skulls;  he  silso  remarks  that  it  more  frequently 
occurs  in  prog-nathous  skulls,  not  of  the  Aryan  race,  than  in  others. 
See  also  G.  Delorenzi  on  the  same  subject;  *Tre  nuovi  casi  d'  anomalia 
deir  osso  malare,'  Torino,  1872.  Also,  E.  Morselli,  'Sopra  una  rara 
anomalia  dell'  osso  malare,'  Modena,  1872.  Still  more  recently  Gruber 
has  written  a  pamphlet  on  the  division  of  this  bone.  I  give  these 
references  because  a  reviewer,  without  any  grounds  or  scruples,  has 
thrown  doubts  on  my  statements. 

^  A  whole  series  of  cases  is  given  by  Isid.  Geoffrey  St.-Hilaire,  'Hist, 
des  Anomalies,'  torn.  iii.  p.  437.  A  reviewer  ('Journal  of  Anat.  and 
Physiolog-y,'  1871,  p.  366)  blames  me  much  for  not  having'  discussed  the 
numerous  cases,  which  have  been  recorded,  of  various  parts  arrested  in 
their  development.  He  says  that,  according-  to  my  theory,  "every  tran- 
"sient  condition  of  an  org^an,  during-  its  development,  is  not  only  a 
"means  to  an  end,  but  once  was  an  end  in  itself."  This  does  not  seem  to 
me  necessarily  to  hold  good.  Why  should  not  variations  occur  during 
an  early  period  of  development,  having  no  relation  to  reversion;  yet 
such  variations  might  be  preserved  and  accumulated,  if  In  any  wa.y  ser- 
viceable, for  instance,  in  shortening  and  simplifying  the  course  of  de* 


MANNER   OF  DEVELOPMENT.  39 

In  man,  the  canine  teeth  are  perfectly  efficient  instruments  for 
mastication.  But  their  true  canine  character,  as  Owen*-  remarks, 
"is  indicated  by  the  conical  form  of  the  crown,  which  terminates 
"in  an  obtuse  point,  is  convex  outward  and  flat  or  sub-concave 
"within,  at  the  base  of  which  surface  there  is  a  feeble  prominence. 
"The  conical  form  is  best  expressed  in  the  Melanian  races,  espe- 
"cially  the  Australian.  The  canine  is  more  deeply  implanted, 
"and  by  a  stronger  fang  than  the  incisors."  Nevertheless,  this 
tooth  no  longer  serves  man  as  a  special  weapon  for  tearing  his 
enemies  or  prey;  it  may,  therefore,  as  far  as  its  proper  function 
is  concerned,  be  considered  as  rudimentary.  In  every  large  col- 
lection of  human  skulls  some  may  be  found,  as  Hackel*^  observes, 
with  the  canine  teeth  projecting  considerably  beyond  the  others 
in  the  same  manner  as  in  the  anthropomorphous  apes,  but  in  a 
less  degree.  In  these  cases,  open  spaces  between  the  teeth  in 
the  one  jaw  are  left  for  the  reception  of  the  canines  of  the  opposite 
jaw.  An  interspace  of  this  kind  in  a  Kaffir  skull,  figured  by  Wag- 
ner, is  surprisingly  wide."  Considering  how  few  are  the  ancient 
skulls  which  have  been  examined,  compared  to  recent  skulls,  it 
is  an  interesting  fact  that  in  at  least  three  cases  the  canines  pro- 
ject largely;  and  in  the  Naulette  jaw  they  are  spoken  of  as  enor- 
mous.*^ 

Of  the  anthropomorphous  apes  the  males  alone  have  their 
canines  fully  developed;  but  in  the  female  gorilla,  and  in  a  less 
degree  in  the  female  orang,  these  teeth  project  considerably  beyond 
the  others;  therefore  the  fact,  of  which  I  have  been  assured, 
that  women  sometimes  have  considerably  projecting  canines,  is 
no  serious  objection  to  the  belief  that  their  occasional  great  de- 
velopment in  man  is  a  case  of  reversion  to  an  ape-like  progenitor. 
He  who  rejects  with  scorn  the  belief  that  the  shape  of  his  own 
canines,  and  their  occasional  great  development  in  other  men, 
are  due  to  our  early  forefathers  having  been  provided  with  these 
formidable  weapons,  will  probably  reveal,  by  sneering,  the  line  of 
his  descent.  For  though  he  no  longer  intends,  nor  has  the  power, 
to  use  these  teeth  as  weapons,  he  will  unconsciously  retract  his 
"snarling  muscles"  (thus  named  by  Sir  C.  Bell),*^  so  as  to  expose 
them  ready  for  action,  like  a  dog  prepared  to  fight. 


velopment?  And  again,  why  should  not  injurious  abnormalities,  such 
as  atrophied  or  hypertrophied  parts,  which  have  no  relation  to  a  for- 
mer state  of  existence,  occur  at  an  early  period,  as  well  as  during- 
maturity? 

*2  'Anatomy  of  Vertebrates,'  vol.  iii.  1868,  p.  323. 

^  'Generelle  Morphologic,'  1886,  B.  ii.  s.  civ. 

**  Carl  Vogt's  'Lectures  on  Man,'  Eng.  translat.  1864,  p.  151. 

*^  C.  Carter  Blake,  on  a  jaw  from  La  Naulette,  'Anthropolog.  Review,' 
1867,  p.  295.    Schaaffhausen,  ibid.  1868,  p.  426. 

*«  'The  Anatomy  of  Expression,'  1844,  pp.  110,  131. 


40  THE  DESCENT  OF  MAN. 

Many  muscles  are  occasionally  developed  in  man,  which  ai-ts 
proper  to  the  Quadrumana  or  other  mammals.  Professor  Vlaco- 
vich^'  examined  forty  male  subjects,  and  found  a  muscle,  called 
by  him  the  ischio-pubic,  in  nineteen  of  them;  in  three  others 
there  was  a  ligament  which  represented  this  muscle;  and  in  the 
remaining  eighteen  no  trace  of  it.  In  only  two  out  of  thirty 
female  subjects  was  this  muscle  developed  on  both  sides,  but  in 
three  others  the  rudimentary  ligament  was  present.  This  muscle, 
therefore,  appears  to  be  much  more  common  in  the  male  than  in 
the  female  sex;  and  on  the  belief  in  the  descent  of  man  from 
some  lower  form,  the  fact  is  intelligible;  for  it  has  been  detected 
in  several  of  the  lower  animals,  and  in  all  of  these  it  serves 
exclusively  to  aid  the  male  in  the  act  of  reproduction. 

Mr.  J.  Wood,  in  his  valuable  series  of  papers,*'  has  minutely 
described  a  vast  number  of  muscular  variations  in  man,  which 
resemble  normal  structures  in  the  lower  animals.  The  muscles 
which  closely  resemble  those  regularly  present  in  our  nearest 
allies,  the  Quadrumana,  are  too  numerous  to  be  here  even  speci- 
fied. In  a  single  male  subject,  having  a  strong  bodily  frame,  and 
well-formed  skull,  no  less  than  seven  muscular  variations  were 
observed,  all  of  which  plainly  represented  muscles  proper  to 
various  kinds  of  apes.  This  man,  for  instance,  had  on  both  sides 
of  his  neck  a  true  and  powerful  "levator  claviculse,"  such  as  is 
found  in  all  kinds  of  apes,  and  which  is  said  to  occur  in  about 
one  out  of  sixty  human  subjects.*^  Again,  this  man  had  "a  special 
"abductor  of  the  metatarsal  bone  of  the  fifth  digit,  such  as  Pro- 
"fessor  Huxley  and  Mr.  Flower  have  shown  to  exist  uniformly 
"in  the  higher  and  lower  apes."  I  will  give  only  two  additional 
cases;  the  acromio-basilar  muscle  is  found  in  all  mammals  below 
man,  and  seems  to  be  correlated  with  a  quadrupedal  gait,^°  and 
it  occurs  in  about  one  out  of  sixty  human  subjects.    In  the  lower 

*'7  Quoted  by  Prof.  Canestrini  in  the  'Annuario,'  &c.,  1867,  p.  90. 

48  These  papers  deserve  careful  study  by  any  one  who  desires  to  learn 
how  frequently  our  muscles  vary,  and  in  varying-  come  to  resemble 
those  of  the  Quadrumana.  The  following  references  relate  to  the  few 
points  touched  on  in  my  text:  'Proc.  Poyal  Soc.  vol.  xiv.  1865,  pp.  379- 
384;  vol.  XV.  1866,  pp.  241,  242;  vol.  xv.  1867,  p.  544;  vol.  xvi.  1868,  p.  524. 
I  may  here  add  that  Dr.  Murie  and  Mr.  St.  George  Mivart  have  shown 
in  their  Memoir  on  the  Lemuroidea  ('Transact.  Zoolog.  Soc'  vol.  vii. 
1869,  p.  96),  how  extraordinarily  variable  some  of  the  muscles  are  in 
these  animals,  the  lowest  members  of  the  Primates.  Gradations,  also, 
in  the  muscles  leading  to  structures  found  in  animals  still  lower  in 
the  scale,  are  numerous  in  the  Lem.uroidea. 

49  See  also  Prof,  Macalister  in  'Proc.  R.  Irish  Academy,'  vol.  x.  1868, 
p.  124. 

K>  Mr.  Champneys  in  'Journal  of  Anat.  and  Phys.'  Nov.,  1871,  p.  178. 


MANNER   OF  DEVELOPMENT.  41 

extremities  Mr.  Bradley^^  found  an  abductor  ossis  metatarsi  quinti 
in  both  feet  of  man;  this  muscle  had  not  up  to  that  time  been 
recorded  in  mankind,  but  is  always  present  in  the  anthropomor- 
phous apes.  The  muscles  of  the  hands  and  arms— parts  of  which 
are  so  eminently  characteristic  of  man — are  extremely  liable  to 
vary,  so  as  to  resemble  the  corresponding  muscles  in  the  lower 
animals.'-  Such  resemblances  are  either  perfect  or  imperfect; 
yet  in  the  latter  case  they  are  manifestly  of  a  transitional  nature. 
Certain  variations  are  more  common  in  man,  and  others  in 
woman,  without  our  being  able  to  assign  any  reason.  Mr.  Wood, 
after  describing  numerous  variations,  makes  the  following  preg- 
nant remark:  "Notable  departures  from  the  ordinary  type  of  the 
"muscular  structures  run  in  grooves  or  directions,  which  must 
"be  taken  to  indicate  some  unknown  factor,  of  much  importance 
"to  a  comprehensive  knowledge  of  general  and  scientific  anat- 
omy."" 

That  this  unknown  factor  is  reversion  to  a  former  state  of 
existence  may  be  admitted  as  in  the  highest  degree  probable.^* 
It  is  quite  incredible  that  a  man  should  through  mere  accident 
abnormally  resemble  certain  apes  in  no  less  than  seven  of  his 
muscles,  if  there  had  been  no  genetic  connection  between  them. 


Bi  'Journal  of  Anat.  and  Phys.'  May,  1872,  p.  421. 

^2  Prof.  Macalister  (ibid.  p.  121)  has  tabulated  his  observations,  and 
finds  that  muscular  abnormalities  are  most  frequent  in  the  fore-arms, 
secondly,  in  the  face,  thirdly,  in  the  foot,  &c. 

S3  The  Rev.  Dr.  Haughton,  after  giving-  ('Proc.  R.  Irish  Academy,' 
June  27,  1864,  p.  715),  a  remarkable  case  of  variation  in  the  human 
flexor  pollicis  longus,  adds,  "This  remarkable  example  shows  that  man 
"may  sometimes  possess  the  arrangement  of  tendons  of  thumb  and 
"fingers  characteristic  of  the  macaque;  but  whether  such  a  case  should 
"be  regarded  as  a  macaque  passing  upwards  into  a  man,  or  a  man 
"passing  downward  into  a  macaque,  or  as  a  congenital  freak  of  nature, 
"I  cannot  undertake  to  say."  It  is  satisfactory  to  hear  so  capable 
an  anatomist,  and  so  embittered  an  opponent  of  evolutionism,  admit- 
ting even  the  possibility  of  either  of  his  first  propositions.  Prof.  Mac- 
alister has  also  described  ('Proc.  R.  Irish  Acad.*  vol.  x.  1864,  p.  138) 
variations  in  the  flexor  pollicis  longus,  remarkable  from  their  rela- 
tions to  the  same  muscle  in  the  Quadrumana. 

^  Since  the  first  edition  of  this  book  appeared,  Mr.  Wood  has  pub- 
lished another  memoir  in  the  'Phil.  Transactions,'  1870,  p.  83,  on  the 
varieties  of  the  muscles  of  the  human  neck,  shoulder,  and  chest.  He 
here  shows  how  extremely  variable  these  muscles  are,  and  how  often 
and  how  closely  the  variations  resemble  the  normal  muscles  of  the 
lower  animals.  He  sums  up  by  remarking,  "It  will  be  enough  for  my 
"purpose  if  I  have  su-cceeded  in  showing  the  more  important  forms 
"which,  when  occurring  as  varieties  in  the  human  subject,  tend  to  ex- 
"hibit  in  a  sufiiciently  marked  manner  what  may  be  considered  as 
"proofs  and  examples  of  the  Darwinian  principle  of  reversion  or  law 
"of  inheritance,  in  this  department  of  anatomical  scienca" 


42  :fHE  DESCENT  OP  MAN. 

On  the  other  hand,  if  man  is  descended  from  some  ape-like  crea- 
ture, no  valid  reason  can  be  assigned  why  certain  muscles  should 
not  suddenly  reappear  after  an  interval  of  many  thousand  genera- 
tions, in  the  same  manner  as  with  horses,  asses,  and  mules,  dark- 
colored  stripes  suddenly  reappear  on  the  legs  and  shoulders,  after 
an  interval  of  hundreds,  or  more  probably  of  thousands  of  genera- 
tions. 

These  various  cases  of  reversion  are  so  closely  related  to  those 
of  rudimentary  organs  given  in  the  first  chapter,  that  many  of 
them  might  have  been  indifferently  introduced  either  there  or 
here.  Thus  a  human  uterus  furnished  with  cornua  may  be  said 
to  represent,  in  a  rudimentary  condition,  the  same  organ  in  its 
normal  state  in  certain  mammals.  Some  parts  which  are  rudi- 
mentary in  man,  as  the  os  coccyx  in  both  sexes,  and  the  mammse 
in  the  male  sex,  are  always  present;  whilst  others,  such  as  the 
supracondyloid  foramen,  only  occasionally  appear,  and  therefore 
might  have  been  introduced  under  the  head  of  reversion.  These 
several  reversionary  structures,  as  well  as  the  strictly  rudi- 
mentary ones,  reveal  the  descent  of  man  from  some  lower  form 
in  an  unmistakable  manner. 

Correlated  Variation. — In  man,  as  in  the  lower  animals,  many 
structures  are  so  intimately  related,  that  when  one  part  varies 
so  does  another,  without  our  being  able,  in  most  cases,  to  assign 
any  reason.  We  cannot  say  whether  the  one  part  governs  the 
other,  or  whether  both  are  governed  by  some  earlier  developed 
part.  Various  monstrosities,  as  I.  Geoffroy  repeatedly  insists,  are 
thus  intimately  connected.  Homologous  structures  are  particu- 
larly liable  to  change  together,  as  we  see  on  the  opposite  sides  of 
the  body,  and  in  the  upper  and  lower  extremities.  Meckel  long 
ago  remarked,  that  when  the  muscles  of  the  arm  depart  from 
their  proper  type,  they  almost  always  imitate  those  of  the  leg; 
and  so,  conversely,  with  the  muscles  of  the  legs.  The  organs  of 
sight  and  hearing,  the  teeth  and  hair,  the  color  of  the  skin  and  of 
the  hair,  color  and  constitution,  are  more  or  less  correlated.^^ 
Professor  Schaaffhausen  first  drew  attention  to  the  relation  ap- 
parently existing  between  a  muscular  frame  and  the  strongly- 
pronounced  supra-orbital  ridges,  which  are  so  characteristic  of 
the  lower  races  of  man. 

Besides  the  variations  which  can  be  grouped  with  more  or 
less  probability  under  the  foregoing  heads,  there  is  a  large  class 
of  variations  which  may  be  provisionally  called  spontaneous,  for 
to  our  ignorance  they  appear  to  arise  without  any  exciting  cause. 
It  can,  however,  be  shown  that  such  variations,  whether  consist- 

Es  The  authorities  for  these  several  statements  are  given  in  my  'Va- 
riation of  Animals  under  Domestication,'  vol.  ii.  pp.  820-335. 


MANNER    OF   DEVELOPMENT.  43 

ing  of  slight  indwidual  differences,  or  of  strongly-marked  and 
abrupt  deviation?  of  structure,  depend  much  more  on  '.he  consti- 
tution of  the  organism  than  on  the  nature  of  the  cov^ditions  to 
which  it  has  been  subjected.^^ 

Rate  of  Increase. — Civilized  populations  have  been  known  under 
favorable  conditions,  as  in  the  United  States,  to  double  their 
numbers  in  twenty-five  years;  and,  according  to  a  calculation  by 
Euler,  this  might  occur  in  a  little  over  twelve  years.^^  At  the 
former  rate,  the  present  population  of  the  United  States  (thirty 
millions),  would  in  657  years  cover  the  whole  terraqueous  globe 
so  thickly,  that  four  men  would  have  to  stand  on  each  square 
yard  of  surface.  The  primary  or  fundamental  check  to  the  con- 
tinued increase  of  man  is  the  difficulty  of  gaining  subsistence,  and 
of  living  in  comfort.  We  may  infer  that  this  is  the  case  from  what 
we  see,  for  instance,  in  the  United  States,  where  subsistence  is 
easy,  and  there  is  plenty  of  room.  If  such  means  were  suddenly 
doubled  in  Great  Britain,  our  number  would  be  quickly  doubled. 
With  civilized  nations  this  primary  check  acts  chiefly  by  restrain- 
ing marriages.  The  greater  death-rate  of  infants  in  the  poorest 
classes  is  also  very  important;  as  well  as  the  greater  mortality, 
from  various  dis'eases,  of  the  inhabitants  of  crowded  and  miserable 
houses,  at  all  ages.  The  effects  of  severe  epidemics  and  wars 
are  soon  counterbalanced,  and  more  than  counterbalanced,  in 
nations  placed  under  favorable  conditions.  Emigration  also  comes 
in  aid  as  a  temporary  check,  but,  with  the  extremely  poor  classes, 
not  to  any  great  extent. 

There  is  reason  to  suspect,  as  Malthus  has  remarked,  that  the 
reproductive  power  is  actually  less  in  barbarous,  than  in  civilized 
races.  We  know  nothing  positively  on  this  head,  for  with  savages 
no  census  has  been  taken;  but  from  the  concurrent  testimony  of 
missionaries,  and  of  others  who  have  long  resided  with  such 
people,  it  appears  that  their  families  are  usually  small,  and 
large  ones  rare.  This  may  be  partly  accounted  for,  as  it  is  be- 
lieved, by  the  women  suckling  their  infants  during  a  long  time; 
but  it  is  highly  probable  that  savages,  who  often  suffer  much 
hardship,  and  who  do  not  obtain  so  much  nutritious  food  as 
civilized  men,  would  be  actually  less  prolific.  I  have  shown  in  a 
former  work,°^  that  all  our  domesticated  quadrupeds  and  birds, 
and  all  our  cultivated  plants,  are  more  fertile  than  the  cor- 
responding species  in  a  state  of  nature.    It  is  no  valid  objection  to 


^  This  whole  subject  has  been  discussed  in  chap,  xxiii.  vol.  ii.  of  my 
'Variation  of  Animals  and  Plants  under  Domestication.' 

"  See  the  ever  memorable  'Essay  on  the  Principle  of  Population,'  by 
the  Rev.  T.  Malthus,  vol.  i.  1826,  p.  6,  517. 

°s  'Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
pp.  111-113,  163. 


44  THE  DESCENT  OF  MAN. 

this  conclusion  tliat  animals  suddenly  supplied  with  an  excess  of 
food,  or  when  grown  very  fat;  and  that  most  plants  on  sudden 
removal  from  very  poor  to  very  rich  soil,  are  rendered  more  or 
less  sterile.  We  might,  therefore,  expect  that  civilized  men, 
who  in  one  sense  are  highly  domesticated,  would  be  more  prolific 
than  wild  men.  It  is  also  probable  that  the  increased  fertility 
of  civilized  nations  would  become,  as  with  our  domestic  animals, 
an  inherited  character:  it  is  at  least  known  that  with  mankind 
a  tendency  to  produce  twins  runs  in  families.^^ 

Notwithstanding  that  savages  appear  to  be  less  prolific  than 
civilized  people,  they  would  no  doubt  rapidly  increase  if  their 
numbers  were  not  by  some  means  rigidly  kept  down.    The  San- 
tali,  or  hill-tribes  of  India,  have  recently  afforded  a  good  illustra- 
tion of  this  fact;    for,  as  shown  by  Mr.  Hunter,*'"  they  have  in- 
creased at  an  extraordinary  rate  since  vaccination  has  been  intro- 
duced, other  pestilences  mitigated,  and  war  sternly  suppressed. 
This  increase,  however,  would  not  have  been  possible  had  not 
these  rude  people  spread  into  the  adjoining  districts,  and  worked 
for  hire.    Savages  almost  always  marry;    yet  there  is  some  pru- 
dential restraint,  for  they  do  not  commonly  marry  at  the  earliest 
possible  age.     The  young  men  are  often  required  to  show  that 
they  can  support  a  wife;    and  they  generally  have  first  to  earn 
the  price  of  which  to  purchase  her  from  her  parents.    With  sav- 
ages the  difficulty  of  obtaining  subsistence  occasionally  limits 
their  number  in  a  much  more  direct  manner  than  with  civilized 
people,  for  all  tribes  periodically  suffer  from  severe  famines.    At 
such  times  savages  are  forced  to  devour  much  bad  food,   and 
their  health  can  hardly  fail  to  be  injured.    Many  accounts  have 
been  published  of  their  protruding  stomachs  and  emaciated  limbs 
after  and  during  famines.     They  are  then,  also,  compelled  to 
wander  much,  and,  as  I  was  assured  in  Australia,  their  infants 
perish  in  large  numbers.     As  famines  are  periodical,  depending 
chiefly  on  extreme  seasons,  all  tribes  must  fluctuate  in  number. 
They  cannot  steadily  and  regularly  increase,  as  there  is  no  arti- 
ficial increase  in  the  supply  of  food.    Savages,  when  hard  pressed, 
encroach  on  each  other's  territories,  and  war  is  the  result;    but 
they  are  indeed  almost  always  at  war  with  their  neighbors.    They 
are  liable  to  many  accidents  on  land  and  water  in  their  search  for 
food;    and  in  some  countries  they  suffer  much  from  the  larger 
beasts  of  prey.    Even  in  India,  districts  have  been  depopulated  by 
the  ravages  of  tigers. 

Malthus  has  discussed  these  several  checks,  but  he  does  not  lay 
stress  enough  on  what  is  probably  the  most  important  of  all, 

^»  Mr,  Sedgwick,  'British  and  Foreign  Medico-Chirurg.  Review,  July, 
1863,  p.  170. 
«o  'The  Annals  of  Rural  Bengal,'  by  W.  W.  Hunter,  1868,  p.  259. 


MANNER    OF   DEVELOPMENT.  45 

namely  infanticide,  especially  of  female  infants,  and  the  habit  of 
procuring  abortion.  These  practices  now  prevail  in  many  quar- 
ters of  the  world;  and  infanticide  seems  formerly  to  have  pre- 
vailed, as  Mr.  M'Lennan"'  has  shown,  on  a  still  more  extensive 
scale.  These  practices  appear  to  have  originated  in  savages  recog- 
nizing the  difficulty,  or  rather  the  impossibility  of  supporting  all 
the  iiifants  that  are  born.  Licentiousness  may  also  be  added  to 
the  foregoing  checks;  but  this  does  not  follow  from  failing  means 
of  subsistence;  though  there  is  reason  to  believe  that  in  some  cases 
(as  in  Japan)  it  has  been  intentionally  encouraged  as  a  means  of 
keeping  down  the  population. 

If  we  look  back  to  an  extremely  remote  epoch,  before  man  had 
arrived  at  the  dignity  of  manhood,  he  would  have  been  guided 
more  by  instinct  and  less  by  reason  than  are  the  lowest  savages 
at  the  present  time.  Our  early  semi-human  progenitors  would  not 
have  practiced  infanticide  or  polyandry;  for  the  instincts  of  the 
lower  animals  are  never  so  perverted^-  as  to  lead  them  regularly  to 
destroy  their  cwn  offspring,  or  to  be  quite  devoid  of  jealousy. 
There  would  have  been  no  prudential  restraint  from  marriage,  and 
the  sexes  would  have  freely  united  at  an  early  age.  Hence  the 
progenitors  of  man  would  have  tended  to  increase  rapidly;  but 
checks  of  some  kind,  either  periodical  or  constant,  must  have 
kept  down  their  numbers,  even  more  severely  than  with  existing 
savages.  What  the  precise  nature  of  these  checks  were,  we  can- 
not say,  any  more  than  with  most  other  animals.  We  know  that 
horses  and  cattle,  which  are  not  extremely  prolific  animals,  when 
first  turned  loose  in  South  America,  increased  at  an  enormous  rate. 
The  elephant,  the  slowest  breeder  of  all  known  animals,  would  in 
a  few  thousand  years  stock  the  whole  world.  The  increase  of 
every  species  of  monkey  must  be  checked  by  some  means;  but  not, 
as  Brehm  remarks,  by  the  attacks  of  beasts  of  prey.  No  one  will 
assume  that  the  actual  power  of  reproduction  in  the  wild  horses 
and  cattle  of  America,  was  at  first  in  any  sensible  degree  in- 

61  'Primitive  Marriage,'  1S65. 

•52  A  writer  in  the  'Spectator'  (March  12th,  1871,  p.  320)  comments  as 
follows  on  this  passag-e:— "Mr.  Darwin  finds  himself  compelled  to  re- 
"introduce  a  new  doctrine  of  the  fall  of  man.  He  shows  that  the 
"instincts  of  the  higher  animals  are  far  nobler  than  the  habits  of  sav- 
"ag-e  races  of  men,  and  he  finds  himself,  therefore,  compelled  to  re- 
"introduce,— in  a  form  of  the  substantial  orthodoxy  of  which  he 
"appears  to  be  quite  unconscious,— and  to  introduce  as  a  scientific 
"hypothesis  the  doctrine  that  man's  gain  of  knowledge  was  the  cause 
"of  a  temporary  but  long-enduring  moral  deterioration  as  indicated  by 
"the  many  foul  customs  especially  as  to  marriage,  of  savage  tribes. 
"What  does  the  Jewish  tradition  of  the  moral  degeneration  of  man 
"through  his  snatching  at  a  knowledge  forbidden  him  by  his  highest 
"instinct  assert  beyond  this?" 


46  THE  DESCENT  OF  MAN. 

creased;  or  that,  as  each  district  became  fully  stocked,  this  same 
power  was  diminished.  No  doubt  in  this  case,  and  in  all  others, 
many  checks  concur,  and  different  checks  under  different  circum- 
stances; periodical  dearths,  depending  on  unfavorable  seasons, 
being  probably  the  most  important  of  all.  So  it  will  have  been 
Y/ith  the  early  progenitors  of  man. 

Natural  Selection.  —We  have  now  seen  that  man  is  variable  in 
body  and  mind;  and  that  the  variations  are  induced,  either  di- 
rectly or  indirectly,  by  the  same  general  causes,  and  obey  the 
same  general  laws,  as  with  the  lower  animals.  Man  has  spread 
widely  over  the  face  of  the  earth,  and  must  have  been  exposed, 
during  his  incessant  migrations,°^  to  the  most  diversified  condi- 
tions. The  inhabitants  of  Tierra  del  Fuego,  the  Cape  of  Good 
Hope,  and  Tasmania  in  the  one  hemisphere,  and  of  the  Arctic 
regions  in  the  other,  must  have  passed  through  many  climates, 
and  changed  their  habits  many  times,  before  they  reached  their 
present  homes.''*  The  early  progenitors  of  man  must  also  have 
tended,  like  all  other  animals,  to  have  increased  beyond  their 
means  of  subsistence;  they  must,  therefore  occasionally  have  been 
exposed  to  a  struggle  for  existence,  and  consequently  to  the  rigid 
law  of  natural  selection.  Beneficial  variations  of  all  kinds  will 
thus,  either  occasionally  or  habitually,  have  been  preserved,  and 
injurious  ones  eliminated.  I  do  not  refer  to  strongly-marked  de- 
viations of  structure,  which  occur  only  at  long  intervals  of  time, 
but  to  mere  individual  differences.  We  know,  for  instance,  that 
the  muscles  of  our  hands  and  feet,  which  determine  our  powers  of 
movement,  are  liable,  like  those  of  the  lower  animals,^  to  incessant 
variability.  If  then  the  progenitors  of  man  inhabiting  any  dis- 
trict, especially  one  undergoing  some  change  in  its  conditions, 
were  divided  into  two  equal  bodies,  the  one  half  which  included  all 
the  individuals  best  adapted  by  their  powers  of  movement  for  gain- 
ing subsistence,  or  for  defending  themselves,  would  on  an  average 
survive  in  greater  numbers,  and  procreate  more  offspring  than  the 
other  and  less  well  endowed  half. 
r"  Man  in  the  rudest  state  in  which  he  now  exists  is  the  most  domi- 
nant animal  that  has  ever  appeared  on  this  earth.  He  has  spread 
I  more  widely  than  any  other  highly  organized  form;  and  all  others 
xhave  yielded  before  him.  He  manifestly  owes  this  immense  superi- 

«3  See  some  g-ood  remarks  to  this  effect  by  W.  Stanley  Jevons,  "A 
"Deduction  from.  Darwin's  Theory,"  'Nature,'  1869,  p.  231. 

«*  Latham,  'Man  and  his  Migrations,'  1851,  p.  135. 

«"  Messrs.  Murie  and  Mivart  in  their  'Anatomy  of  the  Lemuroidea' 
('Transact.  Zoolog.  Soc.  vol.  vii.  1869,  pp.  96-98)  say,  "some  muscles  are 
"so  irregular  in  their  distribution  that  they  cannot  be  well  classed  in 
"any  of  the  above  groups."  These  muscles  differ  even  on  the  oppo- 
site sides  of  the  same  individual. 


MANNER   OF  DEVELOPMENT.  47 

ority  to  his  intellectual  faculties,  to  his  social  habits,  which  lead 
him  to  aid  and  defend  his  fellows,  and  to  his  corporeal  structure. 
The  supreme  importance  of  these  characters  has  been  proved  by 
the  final  arbitrament  of  the  battle  for  life.  Through  his  powers 
of  intellect,  articulate  language  has  been  evolved;  and  on  this  his 
wonderful  advancement  has  mainly  depended.  As  Mr.  Chauncey 
Wright  remarks:®^  "a  psychological  analysis  of  the  faculty  of  lan- 
"guage  shows,  that  even  the  smallest  proficiency  in  it  might  re- 
"quire  more  brain  power  than  the  greatest  proficiency  in  any  other 
"direction."  He  has  invented  and  is  able  to  use  various  weapons, 
tools,  traps,  &c.,  with  which  he  defends  himself,  kills  or  catches 
prey,  and  otherwise  obtains  food.  He  has  made  rafts  or  canoes 
for  fishing  or  crossing  over  to  neighboring  fertile  islands.  He 
has  discovered  the  art  of  making  fire,  by  which  hard  and  stringy 
roots  can  be  rendered  digestible,  and  poisonous  roots  or  herbs 
innocuous.  This  discovery  of  fire,  probably  the  greatest  ever 
made  by  man,  excepting  language,  dates  from  before  the  dawn  of 
history.  These  several  inventions,  by  which  man  in  the  rudest 
state  has  become  so  preeminent,  are  the  direct  results  of  the  devel- 
opment of  his  powers  of  observation,  memory,  curiosity,  imagina- 
tion, and  reason.  I  cannot,  therefore,  understand  how  it  is  that 
Mr.  Wallace"  maintains,  that  "natural  selection  could  only  have 
"endowed  the  savage  with  a  brain  a  little  superior  to  that  of  an 
"ape." 

Although  the  intellectual  powers  and  social  habits  of  man  are 
of  paramount  importance  to  him,  we  must  not  underrate  the  im- 
portance of  his  bodily  structure,  to  which  subject  the  remainder  of 
this  chapter  will  be  devoted;  the  development  of  the  intellectual 
and  social  or  moral  faculties  being  discussed  in  a  later  chapter. 

Even  to  hammer  with  precision  is  no  easy  matter,  as  every  one 


«8  Limits  of  Natural  Selection,  'North  American  Review,'  Oct,  1870, 
p.  295. 

67  'Quarterly  Review,'  AprO,  1869,  p.  392.  This  subject  is  more  fully 
discussed  in  Mr.  Wallace's  'Contributions  to  the  Theory  of  Natural 
Selection,'  1870,  in  which  all  the  essays  referred  to  in  this  work  are 
republished.  The  'Essay  on  Man'  has  been  ably  criticized  by  Prof. 
Claparede,  one  of  the  most  disting-uished  zoologists  in  Europe,  in  an 
article  published  in  the  'Bibliotheque  Universelle,*  June,  1870.  The  re- 
mark quoted  in  my  text  will  surprise  every  one  who  has  read  Mr. 
Wallace's  celebrated  paper*  on  'The  Origin  of  Human  Races  deduced 
from  the  Theory  of  Natural  Selection,'  originally  published  in  the  'An- 
thropological Review,'  May,  1864,  p.  clviii.  I  cannot  here  resist  quot- 
ing a  most  just  remark  by  Sir  J.  Lubbock  ('Prehistoric  Times,'  18G5, 
p.  479)  in  reference  to  this  paper,  namely,  that  Mr.  Wallace,  "with 
"characteristic  unselfishness,  ascribes  it  (i.  e.  the  idea  of  natural  selec- 
"tion)  unreservedly  to  Mr.  Darwin,  althoug-h,  as  is  well  known,  he 
"struck  out  the  idea  independently,  and  published  it,  though  not  with 
"the  same  elaboration,  at  the  same  time." 


4g  THE  DESCENT  OF  MAN. 

who  has  tried  to  learn  carpentry  will  admit.  To  throw  a  stone 
with  as  true  an  aim  as  a  Fuegian  in  defending  himself,  or  in  kill- 
ing birds,  requires  the  most  consummate  perfection  in  the  cor- 
related action  of  the  muscles  of  the  hand,  arm,  and  shoulder,  and, 
further,  a  fine  sense  of  touch.  In  throwing  a  stone  or  spear,  and 
in  many  other  actions,  a  man  must  stand  firmly  on  his  feet,  and 
this  again  demands  the  perfect  co-adaptation  of  numerous  mus- 
cles. To  chip  a  flint  into  the  rudest  tool,  or  to  form  a  barbed  spear 
or  hook  from  a  bone,  demands  the  use  of  a  perfect  hand;  for,  as 
a  most  capable  judge,  Mr.  Schoolcraft,''^  remarks,  the  shaping  frag- 
ments of  stone  into  knives,  lances,  or  arrow-heads,  shows  "ex- 
traordinary ability  and  long  practice."  This  is  to  a  great  extent 
proved  by  the  fact  that  primeval  men  practiced  a  division  of  labor; 
each  man  did  not  manufacture  his  own  flint  tools  or  rude  pottery, 
but  certain  individuals  appear  to  have  devoted  themselves  to  such 
work,  no  doubt  receiving  in  exchange  the  produce  of  the  chase. 
Archgeologists  are  convinced  that  an  enormous  interval  of  time 
elapsed  before  our  ancestors  thought  of  grinding  chipped  flints 
into  smooth  tools.  One  can  hardly  doubt,  that  a  man-like  animal 
who  possessed  a  hand  and  arm  sufficiently  perfect  to  throw  a  stone 
with  precision,  or  to  form  a  flint  into  a  rude  tool,  could,  with  suf- 
ficient practice,  as  far  as  mechanical  skill  alone  is  concerned* 
make  almost  anything  which  a  civilized  man  can  make.  The 
structure  of  the  hand  in  this  respect  may  be  compared  with  that  of 
the  vocal  organs,  which  in  the  apes  are  used  for  uttering  various 
signal-cries,  or,  as  in  one  genus,  musical  cadences;  but  in  man  the 
closely  similar  vocal  organs  have  become  adapted  through  the  in- 
herited effects  of  use  for  the  utterance  of  articulate  language. 

Turning  now  to  the  nearest  allies  of  men,  and  therefore  to  the 
best  representatives  of  our  early  progenitors,  we  find  that  the 
hands  of  the  Quadrumana  are  constructed  on  the  same  general 
pattern  as  our  own,  but  are  far  less  perfectly  adapted  for  diver- 
sified uses.  Their  hands  do  not  serve  for  locomotion  so  well  as 
the  feet  of  a  dog;  as  may  be  seen  in  such  monkeys  as  the  chim- 
panzee and  orang,  which  walk  on  the  outer  margins  of  the  palms, 
or  on  the  knuckles.''^  Their  hands,  however,  are  admirably  adapt- 
ed for  climbing  trees.  Monkeys  seize  thin  branches  or  ropes, 
with  the  thumb  on  one  side  and  the  fingers  and  palm  on  the  other, 
in  the  same  manner  as  we  do.  They  can  thus  also  lift  rather  large 
objects,  such  as  the  neck  of  a  bottle  to  their  mouths.  Baboons 
turn  over  stones,  and  scratch  up  roots  with  their  hands.  They 
seize  nuts,  insects,  or  other  small  objects  with  the  thumb  in  oppo- 


«8  Quoted  by  Mr.  Lawson  Tait  in  his  'Law  of  Natural  Selection,'— 
•Dublin  Quarterly  Journal  of  Medical  Science,'  Feb.  1869.  Dr.  Keller 
Is  likewise  quoted  to  the  same  effect. 

^^  Owen,  'Anatomy  of  Vertebrates,'  vol,  iii.  p.  71. 


MANNER   OF  DEVELOPME2NT.  49 

sition  to  the  fingers,  and  no  doubt  they  thus  extract  eggs  and  the 
young  from  the  nests  of  birds.  American  monkej^s  beat  the  wild 
oranges  on  the  branches  until  the  rind  is  cracked,  and  then  tear  it 
off  with  the  fingers  of  the  two  hands.  In  a  wild  state  they  break 
open  hard  fruits  with  stones.  Other  monkeys  open  mussel-shells 
with  the  two  thumbs.  With  their  fingers  they  pull  out  thorns  and 
burs,  and  hunt  for  each  other's  parasites.  They  roll  down  stones, 
or  throw  them  at  their  enemies:  nevertheless,  they  are  clumsy  in 
these  various  actions,  and,  as  I  have  myself  seen,  are  quite  unable 
to  throw  a  stone  with  precision. 

It  seems  to  me  far  from  true  that  because  "objects  are  grasped 
'^clumsily"  by  monkeys,  "a  much  less  specialized  organ  of  pre- 
"hension"  would  have  served  them'°  equally  well  with  their  pres- 
ent hands.  On  the  contrary,  I  see  no  reason  to  doubt  that  more 
perfectly  constructed  hands  would  have  been  an  advantage  to 
them,  provided  that  they  were  not  thus  rendered  less  fitted  for 
climbirtg  trees.  We  may  suspect  that  a  hand  as  perfect  as  that 
oi  man  would  have  been  disadvantageous  for  climbing;  for  the 
most  arboreal  monkeys  in  the  world,  namely,  Ateles  in  America, 
Colobus  in  Africa,  and  Hylobates  in  Asia,  are  either  thumbless,  or 
their  toes  partially  cohere,  so  that  their  limbs  are  converted  into 
mere  grasping  hooks.^^ 

As  soon  as  some  ancient  member  in  the  great  series  of  the 
Primates  came  to  be  less  arboreal,  owing  to  a  change  in  its  man- 
ner of  procuring  subsistence,  or  to  some  change  in  the  surround- 
ing conditions,  its  habitual  manner  of  progression  would  have  been 
modified r  and  thus  it  would  have  been  rendered  more  strictly 
quadrupedal  or  bipedal.  Baboons  frequent  hilly  and  rocky  dis- 
tricts, and  only  from  necessity  climb  high  trees  ;^2  and  they  have 
acquired  almost  the  gait  of  a  dog.  Man  alone  has  become  a  biped; 
and  we  can,  I  think,  partly  see  how  he  has  come  to  assume  his 
erect  attitude,  which  forms  one  of  his  most  conspicuous  charac- 
ters. Man  could  not  have  attained  his  present  dominant  position 
in  the  world  without  the  use  of  his  hands,  which  are  so  admirably 
adapted  to  act  in  obedience  to  his  will.  Sir  C.  BelF^  insists  that 
"the  hand  supplies  all  instruments,  and  by  its  correspondence  with 
"the  intellect  gives  him  universal  dominion."    But  the  hands  and 


■^0  'Quarterly  Review,'  April,  1869,  p.  392. 

■^1  In  Hylobates  syndactylus,  as  the  name  expresses,  two  of  the  toes 
reg-ularly  cohere;  and  this,  as  Mr.  Blyth  informs  me  is  occasionally 
the  case  with  the  toes  of  H.  agilis,  lar,  and  leuciscus.  Colobus  is 
strictly  arboreal  and  extraordinarily  active  (Brehm,  'Thierleben,'  B.  1, 
s.  50),  but  whether  a  better  climber  than  the  species  of  the  allied  gen- 
era, I  do  not  know.  It  deserves  notice  that  the  feet  of  the  sloths,  the 
most  arboreal  animals  in  the  world,  are  wonderfully  hook-like. 

'2  Brehm,  'Thierleben,'  B.  i.  s.  80. 

'3  "The  Hand,"  &c.  'Bridgewater  Treatise/  1833,  p.  38. 
5 


50  THE  DESCENT  OF  MAN. 

arms  could  hardly  have  become  perfect  enough  to  have  manu- 
factured weapons,  or  to  have  hurled  stones  and  spears  with  a  true 
aim,  as  long  as  they  were  habitually  used  for  locomotion  and  for 
supporting  the  whole  weight  of  the  body,  or,  as  before  remarked, 
so  long  as  they  were  especially  fitted  for  climbing  trees.  Such 
rough  treatment  would  also  have  blunted  the  sense  of  touch,  on 
which  their  delicate  use  largely  depends.  From  these  causes 
alone  it  would  have  been  an  advantage  to  man  to  become  a  biped; 
but  for  many  actions  it  is  indispensable  that  the  arms  and  whole 
upper  part  of  the  body  should  be  free;  and  h©  must  for  this  end 
stand  firmly  on  his  feet.  To  gain  this  great  advantage,  the  feet 
have  been  rendered  fiat;  and  the  great  toe  has  been  peculiarly 
modified,  though  this  has  entailed  the  almost  complete  loss  of  its 
power  of  prehension.  It  accords  with  the  principle  of  the  division 
of  physiological  labor,  prevailing  throughout  the  animal  king- 
dom, that  as  the  hands  became  perfected  for  prehension,  the  feet 
should  have  become  perfected  for  support  and  locomotion.  With 
some  savages,  however,  the  foot  has  not  altogether  lost  its  pre- 
hensile power,  as  shown  by  their  manner  of  climbing  trees,  and  of 
using  them  in  other  ways.^^ 

If  it  be  an  advantage  to  man  to  stand  firmly  on  his  feet  and 
to  have  his  hands  and  arms  free,  of  which,  from  his  pre-emi« 
nent  success  in  the  battle  of  life,  there  can  be  no  doubt,  then  I 
can  see  no  reason  why  it  should  not  have  been  advantageous 
to  the  progenitors  of  man  to  have  become  more  and  more 
erect  or  bipedal.  They  would  thus  have  been  better  able  to  de- 
fend themselves  with  stones  or  clubs,  to  attack  their  prey, 
or  otherwise  to  obtain  food.  The  best  built  individuals 
would  in  the  long  run  have  succeeded  best,  and  have  sur- 
vived in  larger  numbers.  If  the  gorilla  and  a  few  allied 
forms  had  become  extinct,  it  might  have  been  argued,  with 
great  force  and  apparent  truth,  that  an  animal  could  not  have 
been  gradually  converted  from  a  quadruped  into  a  biped,  as  all 
the  individuals  in  an  intermediate  condition  would  have  been 
miserably  ill-fitted  for  progression.  But  we  know  (and  this  is  well 
worthy  of  refiection)  that  the  anthropomorphous  apes  are  now 
actually  in  an  intermediate  condition;  and  no  one  doubts  that 
they  are  on  the  whole  well  adapted  for  their  conditions  of  life. 
Thus  the  gorilla  runs  with  a  sidelong  shambling  gait,  but  more 

'*  Hackel  has  an  excellent  discussion  on  the  steps  by  which  man  be- 
came a  biped:  'Naturliche  Schopfung-sgeschichte,'  1868,  s.  507,  Dr. 
Buchner  ('Conferences  sur  la  Theorie  Darwinienne,'  1869,  p.  135)  has 
given  good  cases  of  the  use  of  the  foot  as  a  prehensile  organ  by  man; 
ajad  has  also  written  on  the  manner  of  progression  of  the  higher  apes, 
lo  which  I  allude  in  the  following  paragraph:  see  also  Owen  ('Anat- 
omy of  Vertebrates,'  vol.  iii.  p.  71)  on  this  latter  subject. 


MANNER   OF  DEVELOPMENT.  51 

commonly  progresses  by  resting  on  its  bent  hands.  The  long- 
armed  apes  occasionally  use  their  arms  like  crutches,  swinging 
their  bodies  forward  between  them,  and  some  kinds  of  Hylobates, 
without  having  been  taught,  can  walk  or  run  upright  with  tolerable 
quickness;  yet  they  move  awkwardly,  and  much  less  securely  than 
man.  We  see,  in  short,  in  existing  monkeys  a  manner  of  progres- 
sion intermediate  between  that  of  a  quadruped  and  a  biped;  but, 
as  an  unprejudiced  judge'^  insists,  the  anthropomorphous  apes 
approach  in  structure  more  nearly  to  the  bipedal  than  to  the  quad- 
rupedal type. 

As  the  progenitors  of  man  became  more  and  more  erect,  with 
their  hands  and  arms  more  and  more  modified  for  prehension  and 
other  purposes,  with  their  feet  and  legs  at  the  same  time  trans- 
formed for  firm  support  and  progression,  endless  other  changes  of 
structure  would  have  become  necessary.  The  pelvis  would  have 
to  be  broadened,  the  spine  peculiarly  curved,  and  the  head  fixed  in 
an  altered  position,  all  which  changes  have  been  attained  by  man. 
Prof.  Schaaffhausen"*'  maintains  that  "the  powerful  mastoid  proc- 
"esses  of  the  human  skull  are  the  result  of  his  erect  position;" 
and  these  processes  are  absent  in  the  orang,  chimpanzee,  &c.,  and 
are  smaller  in  the  gorilla  than  in  man.  Various  other  structures, 
which  appear  connected  with  man's  erect  position,  might  here 
have  been  added.  It  is  very  difficult  to  decide  how  far  these  cor- 
related modifications  are  the  result  of  natural  selection,  and  how 
far  of  the  inherited  effects  of  the  increased  use  of  certain  parts,  or 
of  the  action  of  one  part  on  another.  No  doubt  these  means  of 
change  often  co-operate:  thus  when  certain  muscles,  and  the 
crests  of  bone  to  which  they  are  attached,  become  enlarged  by  ha- 
bitual use,  this  shows  that  certain  actions  are  habitually  per- 
formed and  must  be  serviceable.  Hence  the  individuals  which 
performed  them  best,  would  tend  to  survive  in  greater  numbers. 

The  free  use  of  the  arms  and  hands,  partly  the  cause  and  partly 
the  result  of  man's  erect  position,  appears  to  have  led  in  an  in- 
direct manner  to  other  modifications  of  structure.  The  early  male 
forefathers  of  man  were,  as  previously  stated,  probably  furnished 
with  great  canine  teeth;  but  as  they  gradually  acquired  the  habit 
of  using  stones,  clubs,  or  other  weapons,  for  fighting  with  their 
enemies  or  rivals,  they  would  use  their  jaws  and  teeth  less  and 
less.  In  this  case,  the  jaws,  together  with  the  teeth,  would  be- 
come reduced  in  size,  as  we  may  feel  almost  sure  from  innumerable 
analogous  cases.    In  a  future  chapter  we  shall  meet  with  a  closely 

■^  Prof .  Broca,  La  Constitution  des  Vertebres  caudales;  'La  Revue 
d'Anthropolog-ie,'  1872,  p.  26  (separate  copy). 

'«  'On  the  Primitive  Form  of  the  Skull,'  translated  in  'Anthropolog- 
ical Review,'  Oct.  1868,  p.  428.  Owen  ('Anatomy  of  Vertebrates,'  vol.  ii. 
1866,  p.  551)  on  the  mastoid  processes  in  the  higher  apes. 


52  THE  DESCENT  OP  MAN. 

parallel  case,  in  the  reduction  or  complete  disappearance  of  the 
canine  teeth  in  male  ruminants,  apparently  in  relation  with  the 
development  of  their  horns;  and  in  horses,  in  relation  to  their 
habit  of  fighting  with  their  incisor  teeth  and  hoofs. 

In  the  adult  male  anthropomorphous  apes,  as  Riitimeyer,^^  and 
others,  have  insisted,  it  is  the  effect  on  the  skull  of  the  great  de- 
velopment of  the  jaw-muscles  that  causes  it  to  differ  so  greatly  in 
many  respects  from  that  of  man,  and  has  given  to  these  animals 
"a  truly  frightful  physiognomy."  Therefore,  as  the  jaws  and 
teeth  in  man's  progenitors  gradually  became  reduced  in  size,  the 
adult  skull  would  have  come  to  resemble  more  and  more  that  of 
existing  man.  As  we  shall  hereafter  see,  a  great  reduction  of  the 
canine  teeth  in  the  males  would  almost  certainly  affect  the  teeth 
of  the  females  through  inheritance. 

As  the  various  mental  faculties  gradually  developed  themselves 
the  brain  would  almost  certainly  become  larger.  No  one,  I  pre- 
sume, doubts  that  the  large  proportion  which  the  size  of  man's 
brain  bears  to  his  body,  compared  to  the  same  proportion  in  the 
gorilla  or  orang,  is  closely  connected  with  his  higher  mental  pow- 
ers. We  meet  with  closely  analogous  facts  with  insects,  for  in 
ants  the  cerebral  ganglia  are  of  extraordinary  dimensions,  and  in 
all  the  Hymenoptera  these  ganglia  are  many  times  larger  than  in 
the  less  intelligent  orders,  such  as  beetles,'^®  On  the  other  hand, 
no  one  supposes  that  the  intelle'ct  of  any  two  animals  or  of  any  two 
men  can  be  accurately  gauged  by  the  cubic  contents  of  their  skulls. 
It  is  certain  that  there  may  be  extraordinary  mental  activity 
with  an  extremely  small  absolute  mass  of  nervous  matter:  thus 
the  wonderfully  diversified  instincts,  mental  powers,  and  affec- 
tions of  ants  are  notorious,  yet  their  cerebral  ganglia  are  not  so 
large  as  the  quarter  of  a  small  pin's  head.  Under  this  point  of 
view,  the  brain  of  an  ant  is  one  of  the  most  marvelous  atoms  of 
matter  in  the  world,  perhaps  more  so  than  the  brain  of  a  man. 

The  belief  that  there  exists  in  man  some  close  relation  between 
the  size  of  the  brain  and  the  development  of  the  intellectual  facul- 
ties is  supported  by  the  comparison  of  the  skulls  of  savage  and  civ- 
ilized races,  of  ancient  and  modern  people,  and  by  the  analogy  of 
the  whole  vertebrate  series.  Dr.  J.  Barnard  Davis  has  proved,^** 
by  many  careful  measurements,  that  the  mean  internal  capacity 
of  the  skull  in  Europeans  is  92.3  cubic  inches;  in  Americans  87.5; 

"  'Die  Grenzen  der  Thierwelt,  eine  Betrachtung  zu  Darwin's  Lehre,' 
1868,  s.  51. 

■^  Dujardin,  'Annalesi  des  Sc.  Nat.,'  3rd  series  Zoolog-.  torn.  xiv.  1850, 
p.  203.  See  also  Mr.  Lowne,  'Anatomy  and  Phys.  of  the  Musca  vomi- 
toria,'  1870,  p.  14.  My  son,  Mr.  F.  Darwin,  dissected  for  me  the  cerebral 
ganglia  of  the  Formica  rufa. 

'»  'Philosophical  Transactions,'  1869,  p.  513. 


MANNER    OF   DEVELOPMENT.  53 

in  Asiatics  87.1;  and  in  Australians  only  81.9  cubic  inches.  Pro- 
fessor Broca*°  found  that  the  nineteenth  century  skulls  from  graves 
in  Paris  were  larger  than  those  from  vaults  of  the  twelfth  century, 
in  the  proportion  of  1484  to  1426;  and  that  the  increased  size,  as 
ascertained  by  measurements,  was  exclusively  in  the  frontal  part 
of  the  skull— the  seat  of  the  intellectual  faculties.  Prichard  is 
persuaded  that  the  present  inhabitants  of  Britain  have  "much 
more  capacious  brain-cases"  than  the  ancient  inhabitants.  Never- 
theless, it  must  be  admitted  that  some  skulls  of  very  high  an- 
tiquity, such  as  the  famous  one  of  Neanderthal,  are  well  developed 
and  capacious.*'  With  respect  to  the  lower  animals,  M.  E.  Lartet,^- 
by  comparing  the  crania  of  tertiary  and  recent  mammals  belong- 
ing to  the  same  groups,  has  come  to  the  remarkable  conclusion 
that  the  brain  is  generally  larger  and  the  convolutions  are  more 
complex  in  the  more  recent  forms.  On  the  other  hand,  I  have 
shown®^  that  the  brains  of  domestic  rabbits  are  considerably  reduc- 
ed in  bulk,  in  comparison  with  those  of  the  wild  rabbit  or  hare; 
and  this  may  be  attributed  to  their  having  been  closely  confined 
during  many  generations,  so  that  they  have  exerted  their  intel- 
lect, instincts,  senses  and  voluntary  movements  but  little. 

The  gradually  increasing  weight  of  the  brain  and  skull  in  man 
must  have  influenced  the  development  of  the  supporting  spinal 
column,  more  especially  whilst  he  was  becoming  erect.  As  this 
change  of  position  was  being  brought  about,  the  internal  pressure 
of  the  brain  will  also  have  influenced  the  form  of  the  skull;  for 
many  facts  show  how  easily  the  skull  is  thus  affected.  Ethnolo- 
gists believe  that  it  is  modified  by  the  kind  of  cradle  in  which  in- 
fants sleep.  Habitual  spasms  of  the  muscles,  and  a  cicatrix  from 
a  severe  burn,  have  permanently  modified  the  facial  bones,  in 
young  persons  whose  heads  have  become  fixed  either  sideways  or 
backwards,  owing  to  disease,  one  of  the  two  eyes  has  changed  its 
position,  and  the  shape  of  the  skull  has  been  altered  apparently  by 

80  'Les  Selections,'  M.  P.  Broca,  'Revue  d'Anthropologies,'  1873;  see 
also,  as  quoted  in  C.  Vogt's  'Lectures  on  Man,'  Eng.  translat.  1864,  pp. 
88,  90.    Prichard,  'Phys.  Hist,  of  Mankind,  vol.  i.  1838,  p.  305. 

81  In  the  interesting-  article  just  referred  to.  Prof.  Broca  has  well 
remarked,  that  in  civilized  nations,  the  average  capacity  of  the  skull 
must  be  lowered  by  the  preservation  of  a  considerable  number  of 
individuals,  weak  in  mind  and  body,  who  would  have  been  promptly 
eliminated  in  the  savage  state.  On  the  other  hand,  with  savages,  the 
average  includes  only  the  more  capable  individuals,  who  have  been 
able  to  survive  under  extremely  hard  conditions  of  life.  Broca  thus 
explains  the  otherwise  inexplicable  fact,  that  the  mean  capacity  of  the 
skull  of  the  ancient  Troglodytes  of  Lozere  is  greater  than  that  of  mod- 
ern Frenchmen. 

82  'Comptes-rendus  des  Sciences,'  &c.,  June  1,  1868. 

83  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
i.  pp.  124-129. 

6 


54  THE  DESCENT  OF  MAN. 

the  pressure  of  the  brain  in  a  new  direction.**  I  have  shown  that 
with  long-eared  rabbits  even  so  trifling  a  cause  as  the  lopping  for- 
ward of  one  ear  drags  forward  almost  every  bone  of  the  skull  on 
that  side;  so  that  the  bones  on  the  opposite  side  no  longer  strictly 
correspond.  Lastly,  if  any  animal  were  to  increase  or  diminisii 
much  in  general  size,  without  any  change  in  its  mental  powers,  or 
if  the  mental  powers  were  to  be  much  increased  or  diminished, 
without  any  great  change  in  the  size  of  the  body,  the  shape  of  the 
skull  would  almost  certainly  be  altered.  I  infer  this  from  my  ob- 
servations on  domestic  rabbits,  some  kinds  of  which  have  become 
very  much  larger  than  the  wild  animal,  whilst  others  have  re- 
tained nearly  the  same  size,  but  in  both  cases  the  brain  has  been 
much  reduced  relatively  to  the  size  of  the  body.  Now  I  was  at  first 
much  surprised  on  finding  that  in  all  these  rabbits  the  skull  had 
become  elongated  or  dolichocephalic;  for  instance,  of  two  skulls 
of  nearly  equal  breadth,  the  one  from  a  wild  rabbit  and  the  other 
from  a  large  domestic  kind,  the  former  was  3.15  and  the  latter 
4.3  inches  in  length.*-^  One  of  the  most  marked  distinctions  in  dif- 
ferent races  of  men  is  that  the  skull  in  some  is  elongated,  and  in 
others  rounded;  and  here  the  explanation  suggested  by  the  case  of 
the  rabbits  may  hold  good;  for  Welcker  finds  that  short  "men  in- 
*'cline  more  to  brachj'^cephaly,  and  tall  men  to  dolichocephaly;"^'' 
and  tall  men  may  be  compared  with  the  larger  and  longer-bodied 
rabbits,  all  of  which  have  elongated  skulls,  or  are  dolichocephalic. 

From  these  several  facts  we  can  understand,  to  a  certain  extent, 
the  means  by  which  the  great  size  and  more  or  less  rounded  form 
of  the  skull  have  been  acquired  by  man;  and  these  are  characters 
eminently  distinctive  of  him  in  comparison  with  the  lower  ani- 
mals. 

Another  most  conspicuous  difference  between  man  and  the  lower 
animals  is  the  nakedness  of  his  skin.  Whales  and  porpoises 
(Cetacea),dugongs  (Sirenia)  and  the  hippopotamus  are  naked;  and 
this  may  be  advantageous  to  them  for  gliding  through  the  water; 
nor  would  it  be  injurious  to  them  from  the  loss  of  warmth,  as 
the  species,  which  inhabit  the  colder  regions,  are  protected  by  a 
thick  layer  of  blubber,  serving  the  same  purpose  as  the  fur  of  seals 

84  Schaaffhausen  ^ves  from  Blumenbach  and  Busch,  the  cases  of  the 
spasms  and  cicatrix,  in  'Anthropolog-.  Review,'  Oct.,  1868,  p.  420.  Dr. 
Jarrold  ('Anthropologia,'  1808,  pp.  115,  116)  adduces  from  Camper  and 
from  his  own  observations,  cases  of  the  modification  of  the  skull  from 
the  head  being  fixed  in  an  unnatural  position.  He  believes  that  in  cer- 
tain trades,  such  as  that  of  a  shoemaker,  where  the  head  is  habitually 
held  forward,  the  forehead  becomes  more  rounded  and  prominent. 

85  'Variation  of  Animals,'  «S:c.,  vol.  i.  p.  117,  on  the  elongation  of  the 
skull;    p.  119,  on  the  effect  of  the  lopping  of  one  ear. 

» Quoted    by    Schaaffhausen,    in    'Anthropolog.    Review,'    Oct.,    1868, 


MANNER  OF  DEVELOPMENT.  55 

and  otters.  Elephants  and  rhinoceroses  are  almost  hairless;  and 
as  certain  extinct  species,  which  formerly  lived  under  an  Arctic 
climate,  were  covered  with  long  wool  or  hair,  it  would  almost  ap- 
pear as  if  the  existing  species  of  both  genera  had  lost  their 
hairy  covering  from  exposure  to  heat.  This  appears  the  more 
probable,  as  the  elephants  in  India  which  live  on  elevated  and 
cool  districts  are  more  hairy^'  than  those  on  the  lowlands.  May 
we  then  infer  that  man  became  divested  of  hair  from  having  abor- 
iginally inhabited  some  tropical  land?  That  the  hair  is  chiefly 
retained  in  the  male  sex  on  the  chest  and  face,  and  in  both  sexes 
at  the  junction  of  all  four  limbs  with  the  trunk,  favors  this  in- 
ference— on  the  assumption  that  the  hair  was  lost  before  man  be- 
came erect;  for  the  parts  which  now  retain  most  hair  would  then 
have  been  most  protected  from  the  heat  of  the  sun.  The  crown 
of  the  head,  however,  offers  a  curious  exception,  for  at  all  times 
it  must  have  been  one  of  the  most  exposed  parts,  yet  it  is  thickly 
clothed  with  hair.  The  fact,  however,  that  the  other  members  of 
the  order  of  Primates,  to  which  man  belongs,  although  inhabiting 
various  hot  regions,  are  well  clothed  with  hair,  generally  thickest 
on  the  upper  surface,^^  is  opposed  to  the  supposition  that  man  be- 
came naked  through  the  action  of  the  sun.  Mr.  Belt  believes*^  that 
within  the  tropics  it  is  an  advantage  to  man  to  be  destitute  of  hair, 
as  he  is  thus  enabled  to  free  himself  of  the  multitude  of  ticks 
(acari)  and  other  parasites,  with  which  he  is  often  infested,  and 
which  sometimes  cause  ulceration.  But  whether  this  evil  is  of 
suflBcient  magnitude  to  have  led  to  the  denudation  of  his  body 
through  natural  selection,  may  be  doubted,  since  none  of  the  many 
quadrupeds  inhabiting  the  tropics  have,  as  far  as  I  know,  ac- 
quired any  specialized  means  of  relief.  The  view  which  seems  to 
me  the  most  probable  it-  that  man,  or  rather  primarily  woman, 
became  divested  of  hair  for  ornamental  purposes,  as  we  shall  see 
under  Sexual  Selection;  and,  according  to  this  belief,  it  is  not  sur- 
prising that  man  should  differ  so  greatly  in  hairiness  from  all 

^  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  619. 

88  Isidore  Geoffrey  St.-Hilaire  remarks  ('Hist.  Nat.  Generale,'  tcm. 
ii.  1859,  pp.  215-217)  on  the  head  of  man  being  covered  with  long  hair; 
also  on  the  upper  vSurfaces  of  monkeys  and  of  other  mammals  being 
more  thickly  clothed  than  the  lower  surfaces.  This  has  likewise  been 
observed  by  various  authors.  Prof.  P.  Gervais  ('Hist.  Nat.  des  Mam- 
miferes,'  torn.  i.  1854,  p.  28),  however,  states  that  in  the  Gorilla  the  hair 
is  thinner  on  the  back,  where  it  is  partly  rubbed  off,  than  on  the 
lower  surface. 

89  The  'Naturalist  in  Nicaragua,'  1874,  p.  209.  As  some  confirmation  of 
Mr.  Belt's  view,  I  may  quote  the  following  passage  from  Sir  W.  Deni- 
son  ('Varieties  of  Vice-Regal  Life'  vol.  i.  1870,  p.  440):  "It  is  said  to  be 
"a  practice  with  the  Australians,  when  the  vermin  get  troublesome, 
"to  singe  themselves." 


56  THE  DESCENT  OF  MAN. 

Other  Primates,  for  characters,  gained  through  sexual  selection, 
often  differ  to  an  extraordinary  degree  in  closely-related  forms. 

According  to  a  popular  impression,  the  absence  of  a  tail  is 
eminently  distinctive  of  man;  but  as  those  apes  which  come  near- 
est to  him  are  destitute  of  this  organ,  its  disappearance  does  not 
relate  exclusively  to  man.  The  tail  often  differs  remarkably  in 
length  within  the  same  genus:  thus  in  some  species  of  Macacus  it 
fs  longer  than  the  whole  body,  and  is  formed  of  twenty-four  verte- 
bra; in  others  it  consists  of  a  scarcely  visible  stump,  containing 
«nly  three  or  four  vertebrae.  In  some  kinds  of  baboons  there  are 
twenty-five,  whilst  in  the  mandrill  there  are  ten  very  small 
stunted  caudal  vertebrae  or,  according  to  Cuvier,^"  sometimes  only 
five.  The  tail,  whether  it  be  long  or  short,  almost  always  tapers 
towards  the  end;  and  this,  I  presume,  results  from  the  atrophy  of 
the  terminal  muscles,  together  with  their  arteries  and  nerves, 
through  disuse,  leading  to  the  atrophy  of  the  terminal  bones.  But 
no  explanation  can  at  present  be  given  of  the  great  diversity  which 
often  occurs  in  its  length.  Here,  however,  we  are  more  specially 
concerned  with  the  complete  external  disappearance  of  the  tail. 
Professor  Broca  has  recently  shown^^  that  the  tail  in  all  quadru- 
peds consists  of  two  portions,  generally  separated  abruptly  from 
each  other;  the  basal  portion  consists  of  vertebrae,  more  or  less 
perfectly  channelled  and  furnished  with  apophyses  like  ordinary 
vertebrae;  whereas  those  of  the  terminal  portion  are  not  chan- 
nelled, are  almost  smooth,  and  scarcely  resemble  true  vertebrae. 
A  tail,  though  not  externally  visible,  is  really  present  in  man  and 
the  anthropomorphous  apes,  and  is  constructed  on  exactly  the 
same  pattern  in  both.  In  the  terminal  portion  the  vertebrae,  con- 
stituting the  OS  coccyx,  are  quite  rudimentary,  being  much  reduced 
in  size  and  number.  In  the  basal  portion,  the  vertebra  are  like- 
wise few,  are  united  firmly  together,  and  are  arrested  in  develop- 
ment; but  they  have  been  rendered  much  broader  and  flatter  than 
the  corresponding  vertebrae  in  the  tails  of  other  animals:  they 
constitute  what  Broca  calls  the  accessory  sacral  vertebrae. 
These  are  of  functional  importance  by  supporting  certain  internal 
parts  and  in  other  ways;  and  their  modification  is  directly  con- 
nected with  the  erect  or  semi-erect  attitude  of  man  and  the  anthro- 
pomorphous apes.  This  conclusion  is  the  more  trustworthy,  as 
Broca  formerly  held  a  different  view,  which  he  has  now  abandoned. 
The  modification,  therefore,  of  the  basal  caudal  vertebrae  in  man 
and  the  higher  apes  may  have  been  effected,  directly  or  indirectly, 
through  natural  selection. 

»o  Mr.  St.  George  Mivart,  'Proc.  Zoolog.  Soc,  1865,  pp.  562,  583.  Dr.  J.  E. 
Gray,  'Cat.  Brit.  Mus. :  Skeletons.'  Owen,  'Anatomy  of  Vertebrates,' 
vol.  ii.  p.  517.    Isidore  Geoff roy,  'Hist.  Nat.  Gen.'  torn.  ii.  p.  244. 

»i  'Revue  d'Anthropologle,'  1872;  'La  Constitution  des  Vertebres  cau- 
dales.' 


MANNER  OF  DEVELOPMENT.  57 

But  what  are  we  to  say  about  the  rudimentary  and  variable 
vertebrae  of  the  terminal  portion  of  the  tail,  forming  the  os 
coccyx?  A  notion  which  has  often  been,  and  will  no  doubt  again 
be  ridiculed,  namely,  that  friction  has  had  something  to  do  with 
the  disappearance  of  the  external  portion  of  the  tail,  is  not  so 
ridiculous  as  it  at  first  appears.  Dr.  Anderson°-  states  that  the 
extremely  short  tail  of  Macacus  brunneus  is  formed  of  eleven  ver- 
tebrae, including  the  imbedded  basal  ones.  The  extremity  is  ten- 
donous  and  contains  no  vertebrae;  this  is  succeeded  by  five  rudi- 
mentary ones,  so  minute  that  together  they  are  only  one  line  and 
a  half  in  length,  and  these  are  permanently  bent  to  one  side  in  the 
shape  of  a  hook.  The  free  part  of  the  tail,  only  a  little  above  an 
inch  in  length,  includes  only  four  more  small  vertebrae.  This 
short  tail  is  carried  erect;  but  about  a  quarter  of  its  total  length  is 
doubled  on  to  itself  to  the  left;  and  this  terminal  part,  which  in- 
cludes the  hook-like  portion,  serves  "to  fill  up  the  interspace  be- 
"tween  the  upper  divergent  portion  of  the  callosities;"  so  that  the 
animal  sits  on  it,  and  thus  renders  it  rough  and  callous.  Dr.  An- 
derson thus  sums  up  his  observations:  "These  facts  seem  to  me 
"to  have  only  one  explanation;  this  tail,  from  its  short  size,  is  in 
"the  monkey's  way  v/hen  it  sits  down,  and  frequently  becomes 
"placed  under  the  animal  while  it  is  in  this  attitude;  and  from  the 
"circumstance  that  it  does  not  extend  beyond  the  extremity  of  the 
"ischial  tuberosities  it  seems  as  if  the  tail  originally  had  been  bent 
"round,  by  the  will  of  the  animal,  into  the  interspace  between  the 
"callosities,  to  escape  being  pressed  between  them  and  the  ground, 
"and  that  in  time  the  curvature  became  permanent,  fitting  in  of 
"itself  when  the  organ  happens  to  be  sat  upon."  Under  these  cir- 
cumstances it  is  not  surprising  that  the  surface  of  the  tail  should 
have  been  roughened  and  rendered  callous;  and  Dr.  Murie,''^  who 
carefully  observed  this  species  in  the  Zoological  Gardens,  as  well 
as  three  other  closely  allied  forms  with  slightly  longer  tails,  says 
that  when  the  animal  sits  down,  the  tail  "is  necessarily  thrust  to 
"one  side  of  the  buttocks;  and  whether  long  or  short  its  root  is 
"consequently  liable  to  be  rubbed  or  chafed."  As  we  now  have 
evidence  that  mutilations  occasionally  produce  an  inherited  ef- 
fect,^^  it  is  not  very  improbable  that  in  short-tailed  monkeys,  the 
projecting  part  of  the  tail,  being  functionally  useless,  should  after 

s2  'Proc.  Zoolog.  Soc.,'  1872,  p.  210. 

«3  'Proc.  Zoolog.  Soc.,'  1872,  p.  786. 

^  I  allude  to  Dr.  Brown-Sequard's  observations  on  the  transmitted  ef- 
fect of  an  operation  causing  epilepsy  in  guinea-pigs,  and  likewise  more 
recently  on  the  analogous  effects  of  cutting  the  sympathetic  nerve  in 
the  neck.  I  shall  hereafter  have  occasion  to  refer  to  Mr.  Salvin's  inter- 
esting case  of  the  apparently  inherited  effects  of  mot-mots  biting  off  the 
barbs  of  their  own  tail-feathers.  See  also  on  the  general  subject  'Vari- 
8.tion  of  Animals  and  Plants  under  Domestication,'  vol.  ii.  pp.  22-24. 


58  THE  DESCENT  OF  MAN. 

man.y  generations  have  become  rudimentary  and  distorted,  from 
being  continually  rubbed  and  chafed.  We  see  the  projecting  part 
in  this  condition  in  the  Macacus  brunneus,  and  absolutely  aborted 
in  the  M.  ecaudatus  an4  in  several  of  the  higher  apes.  Finally, 
then,  as  far  as  we  can  judge,  the  tail  has  disappeared  in  man  and 
the  anthropomorphous  apes,  owing  to  the  terminal  portion  having 
been  injured  by  friction  during  a  long  lapse  of  time;  the  basal 
and  embedded  portion  having  been  reduced  and  modified,  so  as  to 
become  suitable  to  the  erect  or  semi-erect  position. 
'*  I  have  now  endeavored  to  show  that  some  of  the  most  dis- 
tinctive characters  of  man  have  in  all  probability  been  acquired, 
either  directly,  or  more  commonly  indirectly,  through  natural  se- 
lection. We  should  bear  in  mind  that  modifications  in  structure 
or  constitution,  which  do  not  serve  to  adapt  an  organism  to  its 
habits  of  life,  to  the  food  which  it  consumes,  or  passively  to  the 
surrounding  conditions,  cannot  have  been  thus  acquired.  We 
must  not,  however,  be  too  confident  in  deciding  what  modifications 
are  of  service  to  each  being:  v/e  should  remember  how  little  we 
know  about  the  use  of  many  parts,  or  what  changes  in  the  blood 
or  tissues  may  serve  to  fit  an  organism  for  a  new  climate  or  new 
kinds  of  food.  Nor  must  we  forget  the  principle  of  correlation, 
by  which,  as  Isidore  Geoffrey  has  shown  in  the  case  of  man,  many 
strange  deviations  of  structure  are  tied  together.  Independently 
of  correlation,  a  change  in  one  part  often  leads,  through  the  in- 
creased or  decreased  use  of  other  parts,  to  other  changes  of  a 
quite  unexpected  nature.  It  is  also  well  to  reflect  on  such  facts,  as 
the  wonderful  growth  of  galls  on  plants  caused  by  the  poison  of 
an  insect,  and  on  the  remarkable  changes  of  color  in  the  plum- 
age of  parrots  when  fed  on  certain  fishes,  or  inoculated  with  the 
poison  of  toads ;^^  for  we  can  thus  see  that  the  fluids  of  the  system, 
if  altered  for  some  special  purpose,  might  induce  other  changes. 
We  should  especially  bear  in  mind  that  modifications  acquired 
and  continually  used  during  past  ages  for  some  useful  purpose, 
would  probably  become  firmly  fixed,  and  might  be  long  inherited. 

Thus  a  large  yet  undefined  extension  may  safely  be  given  to  the 
direct  and  indirect  results  of  natural  selection;  but  I  now  admit, 
after  reading  the  essay  by  Nageli  on  plants,  and  the  remarks  by 
various  authors  with  respect  to  animals,  more  especially  those  re- 
cently made  by  Professor  Broca,  that  in  the  earlier  editions  of  my 
'Origin  of  Species'  I  perhaps  attributed  too  much  to  the  action  of 
natural  selection  or  the  survival  of  the  fittest.  I  have  altered  the 
fifth  edition  of  the  'Origin'  so  as  to  confine  my  remarks  to  adaptive 
changes  of  structure;  but  I  am  convinced,  from  the  light  gained 
during  even  the  last  few  years,  that  very  many  structures  which 

»5  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
pp.  280,  282. 


MANNiCR  OF  DEVELOPMENT.  59 

now  appear  to  us  useless,  will  hereafter  be  proved  to  be  use- 
ful, and  will  therefore  come  within  the  range  of  natural  selection. 
Nevertheless,  I  did  not  formerly  consider  sufficiently  the  existence 
of  structures,  which,  as  far  as  we  can  at  present  judge,  are  neither 
beneficial  nor  injurious;  and  this  I  believe  to  be  one  of  the  greatest 
oversights  as  yet  detected  in  my  work.  I  may  be  permitted  to  say, 
as  some  excuse,  that  I  had  two  distinct  objects  in  view;  firstly,  to 
show  that  species  had  not  been  separately  created,  and  secondly, 
that  natural  selection  had  been  the  chief  agent  of  change,  though 
largely  aided  by  the  inherited  effects  of  habit,  and  slightly  by  the 
direct  action  of  the  surrounding  conditions.  I  was  not,  however, 
able  to  annul  the  influence  of  my  former  belief,  then  almost  uni- 
versal, that  each  species  had  been  purposely  created;  and  this  led 
to  my  tacit  assumption  that  every  detail  of  structure,  excepting 
rudiments,  was  of  some  special,  though  unrecognized,  service. 
Any  one  with  this  assumption  in  his  mind  would  naturally  ex- 
tend too  far  the  action  of  natural  selection,  either  during  past  or 
present  times.  Some  of  those  who  admit  the  principle  of  evolu- 
tion, but  reject  natural  selection,  seem  to  forget,  when  criticizing 
my  book,  that  I  had  the  above  two  objects  in  view;  hence  if  I  have 
erred  in  giving  to  natural  selection  great  power,  which  I  am  very 
far  from  admitting,  or  in  having  exaggerated  its  power,  which  is 
in  itself  probable,  I  have  at  least,  as  I  hope,  done  good  service  in 
aiding  to  overthrow  the  dogma  of  separate  creations. 

It  is,  as  I  can  now  see,  probable  that  all  organic  beings,  includ- 
ing man,  possess  peculiarities  of  structure,  which  neither  are  now, 
nor  were  formerly  of  any  service  to  them,  and  which,  therefore, 
are  of  no  physiological  importance.  We  know  not  what  produces 
the  numberless  slight  differences  between  the  individuals  of  each 
species,  for  reversion  only  carries  the  problem  a  few  steps  back- 
wards; but  each  peculiarity  must  have  had  its  efficient  cause.  If 
these  causes,  whatever  they  may  be,  were  to  act  more  uniformly 
and  energetically  during  a  lengthened  period  (and  against  this  no 
reason  can  be  assigned),  the  result  would  probably  be  not  a  mere 
slight  individual  difference,  but  a  well-marked  and  constant  modi- 
fication, though  one  of  no  physiological  importance.  Changed 
structures,  which  are  in  no  way  beneficial,  cannot  be  kept  uniform 
through  natural  selection,  though  the  injurious  will  be  thus  elimi- 
nated. Uniformity  of  character  would,  however,  naturally  follow 
from  the  assumed  uniformity  of  the  exciting  causes,  and  likewise 
from  the  free  intercrossing  of  many  individuals.  During  succes- 
sive periods,  the  same  organism  might  in  this  manner  acquire 
successive  modifications,  which  would  be  transmitted  in  a  nearly 
uniform  state  as  long  as  the  exciting  causes  remained  the  same 
and  there  was  free  intercrossing.  Vvlth  respect  to  the  exciting 
causes  we  can  only  say,  as  when  speaking  of  so-called  spontaneous 
variations,  that  they  relate  much  more  closely  to  the  constitution 


60  THE  DESCENT  OF  MAN. 

of  the  varying  organism,  than  to  the  nature  of  the  conditions  to 
which  it  has  been  subjected. 

Conclusion.  — In  this  chapter  we  have  seen  that  as  man  at  the 
present  day  is  liable,  like  every  other  animal,  to  multiform  in- 
dividual differences  or  slight  variations,  so  no  doubt  were  the 
early  progenitors  of  man;  the  variations  being  formerly  induced 
by  the  same  general  causes,  and  governed  by  the  same  general  and 
complex  laws  as  at  present.  As  all  animals  tend  to  multiply  be- 
yond their  means  of  subsistence,  so  it  must  have  been  with  the 
progenitors  of  man;  and  this  would  inevitably  lead  to  a  struggle 
for  existence  and  to  natural  selection.  The  latter  process  would 
be  greatly  aided  by  the  inherited  effects  of  the  increased  use  of 
parts,  and  these  two  processes  would  incessantly  react  on  each 
other.  It  appears,  also,  as  we  shall  hereafter  see,  that  various 
unimportant  characters  have  been  acquired  by  man  through  sex- 
ual selection.  An  unexplained  residuum  of  change  must  be  left  to 
the  assumed  uniform  action  of  those  unknown  agencies,  which 
occasionally  induce  strongly  marked  and  abrupt  deviations  of 
structure  in  our  domestic  productions. 

Judging  from  the  habits  of  savages  and  of  the  greater  number 
of  the  Quadrumana,  primeval  men,  and  even  their  ape-like  pro- 
genitors, probably  lived  in  society.  With  strictly  social  animals, 
natural  selection  sometimes  acts  on  the  individual,  through  the 
preservation  of  variations  which  are  beneficial  to  the  community. 
A  community  which  includes  a  large  number  of  well-endowed  in- 
dividuals increases  in  number,  and  is  victorious  over  other  less 
favored  ones;  even  although  each  separate  member  gains  no  ad- 
vantage over  the  others  of  the  same  community.  Associated  in- 
sects have  thus  acquired  many  remarkable  structures,  which  are 
of  little  or  no  service  to  the  individual,  such  as  the  pollen-collect- 
ing apparatus,  or  the  sting  of  the  worker-bee,  or  the  great  jaws  of 
soldier-ants.  With  the  higher  social  animals,  I  am  not  aware  that 
any  structure  has  been  modified  solely  for  the  good  of  the  com- 
munity, though  some  are  of  secondary  service  to  it.  For  instance, 
the  horns  of  ruminants  and  the  great  canine  teeth  of  baboons  ap- 
pear to  have  been  acquired  by  the  males  as  weapons  for  sexual 
strife,  but  they  are  used  in  defense  of  the  herd  or  troop.  In  regard 
to  certain  mental  powers  the  case,  as  we  shall  see  in  the  fifth 
chapter,  is  wholly  different;  for  these  faculties  have  been  chiefly, 
or  even  exclusively,  gained  for  the  benefit  of  the  community,  and 
the  individuals  thereof,  have  at  the  same  time  gained  an  ad- 
vantage indirectly. 

It  has  often  been  objected  to  such  views  as  the  foregoing,  that 
man  is  one  of  the  most  helpless  and  defenseless  creatures  in  the 
world;  and  that  during  his  early  and  less  well-developed  con- 
dition he  would  have  been   still  more  helpless.    The  Duke  of 


MANNER  OF  DEVELOPMENT.  61 

Argyll,  for  instance,  insists^'  that  "the  human  frame  has  diverged 
"from  the  structure  of  brutes,  in  the  direction  of  greater  physical 
"helplessness  and  weakness.  That  is  to  say,  it  is  a  divergence 
"which  of  all  others  it  is  most  impossible  to  ascribe  to  mere 
"natural  selection."  He  adduces  the  naked  and  unprotected  state 
of  the  body,  the  absence  of  great  teeth  or  claws  for  defense,  the 
small  strength  and  speed  of  man,  and  his  slight  power  of  discover- 
ing food  or  of  avoiding  danger  by  smell.  To  these  deficiencies 
there  might  be  added  one  still  more  serious,  namely,  that  he  can- 
not climb  quickly,  and  so  escape  from  enemies.  The  loss  of  hair 
would  not  have  been  a  great  injury  to  the  inhabitants  of  a  warm 
country.  For  we  know  that  the  unclothed  Fuegians  can  exist  un- 
der a  wretched  climate.  When  we  compare  the  defenseless  state 
of  man  with  that  of  apes,  we  must  remember  that  the  great  canine 
teeth  with  which  the  latter  are  provided,  are  possessed  in  their  full 
development  by  the  males  alone,  and  are  chiefly  used  by  them  for 
fighting  with  their  rivals;  yet  the  females,  which  are  not  thus  pro- 
vided, manage  to  survive. 

In  regard  to  bodily  size  or  strength,  we  do  not  know  whether 
man  is  descended  from  some  small  species,  like  the  chimpanzee, 
or  from  one  as  powerful  as  the  gorilla;  and,  therefore,  we  cannot 
say  whether  man  has  become  larger  and  stronger,  or  smaller  and 
weaker,  than  his  ancestors.  We  should,  however,  bear  in  mind 
that  an  animal  possessing  great  size,  strength,  and  ferocity  and 
which,  like  the  gorilla,  could  defend  itself  from  all  enemies,  would 
not  perhaps  have  become  social;  and  this  would  most  effectually 
have  checked  the  acquirement  of  the  higher  mental  qualities,  such 
as  sympathy  and  the  love  of  his  fellows.  Hence  it  might  have 
been  an  immense  advantage  to  man  to  have  sprung  from  some 
comparatively  weak  creature. 

The  small  strength  and  speed  of  man,  his  want  of  natural  weap- 
ons, &c.,  are  more  than  counterbalanced,  firstly,  by  his  intel- 
lectual powers,  through  which  he  has  formed  for  himself  weap- 
ons, tools,  &c.,  though  still  remaining  in  a  barbarous  state,  and, 
secondly,  by  his  social  qualities  which  lead  him  to  give  and  re- 
ceive aid  from  his  fellow-men.  No  country  in  the  world  abounds 
in  a  greater  degree  with  dangerous  beasts  than  Southern  Africa; 
no  country  presents  more  fearful  physical  hardships  than  the  Arc- 
tic regions;  yet  one  of  the  puniest  of  races,  that  of  the  Bushmen, 
maintains  itself  in  Southern  Africa,  as  do  the  dwarfed  Esquimaux 
in  the  Arctic  regions.  The  ancestors  of  man  were,  no  doubt,  in- 
ferior in  intellect,  and  probably  in  social  disposition,  to  the  lowest 
existing  savages;  but  it  is  quite  conceivable  that  they  might  have 
existed,   or  even  flourished,  if  they  had  advanced  in  intellect, 

»«  'Primeval  Man,'  1869,  p.  66. 


62  THE  DESCENT  OF  MAN. 

v.hilst  gradually  losing  their  brute-like  powers,  such  as  that  of 
climbing  trees,  &c.  But  these  ancestors  would  not  have  been  ex- 
posed to  any  special  danger,  even  if  far  more  helpless  and  defense- 
less than  any  existing  savages,  had  they  inhabited  some  warm  con- 
tinent or  large  island,  such  as  Australia,  New  Guinea,  or  Borneo, 
which  is  now  the  home  of  the  orang.  And  natural  selection  aris- 
ing from  the  competition  of  tribe  with  tribe,  in  some  such  large 
area  as  one  of  these,  together  with  the  inherited  effects  of  habit, 
would,  under  favorable  conditions  have  suflaced  to  raise  man  to 
his  present  high  position  in  the  organic  scale. 


MENTAL    POWERS.  63 


CHAPTER  III. 

COMPARISON  OF  THE  MENTAL  POWERS  OF  MAN  AND 
THE  LOWER  ANIMALS. 

The  difference  in  mental  power  between  the  highest  ape  and  the  lowest 
savage,  immense — Certain  instincts  in  common — The  emotions- 
Curiosity— Imitation— Attention  —  Memory  —  Imagination— P.eason— 
Progressive  improvement— Tools  and  weapons  used  by  animals— Ab- 
straction, self-consciousness— Language— Sense  of  beauty— Belief  in 
God,  spiritual  agencies,  superstitions. 

We  have  seen  in  the  last  two  chapters  that  man  bears  in  his 
bodily  structure  clear  traces  of  his  descent  from  some  lower 
form;  but  it  may  be  urged  that,  as  man  differs  so  greatly  in 
his  mental  power  from  all  other  animals,  there  must  be  some 
error  in  this  conclusion.  No  doubt  the  difference  in  this  respect 
is  enormous,  even  if  we  compare  the  mind  of  one  of  the  lowest 
savages,  who  has  no  words  to  express  any  number  higher  than 
four,  and  who  uses  hardly  any  abstract  terms  for  common  objects 
or  for  the  affections,^  with  that  of  the  most  highly  organized  ape. 
The  difference  would,  no  doubt,  still  remain  immense,  even  if  one 
of  the  higher  apes  had  been  improved  or  civilized  as  much  as 
a  dog  has  been  in  comparison  with  its  parent-form,  the  wolf  or 
jackal.  The  Puegians  rank  amongst  the  lowest  barbarians,  but 
I  was  continually  struck  with  surprise  how  closely  the  three 
natives  on  board  H.M.S.  "Beagle,"  who  had  lived  some  years  in 
England  and  could  talk  a  little  English,  resembled  us  in  disposi- 
tion and  in  most  of  our  mental  faculties.  If  no  organic  being  ex- 
cepting man  had  possessed  any  mental  power,  or  if  his  powers  had 
been  of  a  wholly  different  nature  from  those  of  the  lov/er  animals, 
then  we  should  never  have  been  able  to  convince  ourselves  that 
our  high  faculties  had  been  gradually  developed.  But  it  can  be 
shown  that  there  is  no  fundamental  difference  of  this  kind.  We 
must  also  admit  that  there  is  a  much  wider  interval  in  mental 
power  between  one  of  the  lowest  fishes,  as  a  lamprey  or  lancelet, 

^  See  the  evidence  on  those  points  as  given  by  Lubbock,  'Prehistoric 
V'imes,'  p.  354,  &a 


64  THE  DESCENT  OF  MAN. 

and  one  of  the  higher  apes,  than  between  an  ape  and  man;  yet 
this  interval  is  filled  up  by  numberless  gradations. 

Nor  is  the  difference  slight  in  moral  disposition  between  a 
barbarian,  such  as  the  man  described  by  the  old  navigator  Byron, 
who  dashed  his  child  on  the  rocks  for  dropping  a  basket  of  sea- 
urchins,  and  a  Howard  or  Clarkson;  and  in  intellect  between  a 
savage  who  uses  hardly  any  abstract  terms,  and  a  Newton  or 
Shakspeare.  Differences  of  this  kind  between  the  highest  men 
of  the  highest  races  and  the  lowest  savages,  are  connected  by  the 
finest  gradations.  Therefore  it  is  possible  that  they  might  pass 
and  be  developed  into  each  other. 

My  object  in  this  chapter  is  to,  show  that  there  is  no  funda- 
mental difference  between  man  and  the  higher  mammals  in  their 
mental  faculties.  Each  division  of  the  subject  might  have  been 
extended  into  a  separate  essay,  but  must  here  be  treated  briefly. 
As  no  classification  of  the  mental  powers  has  been  universally 
accepted,  I  shall  arrange  my  remarks  in  the  order  most  con- 
venient for  my  purpose;  and  will  select  those  facts  which  have 
struck  me  most,  with  the  hope  that  they  may  produce  some 
effect  on  the  reader. 

With  respect  to  animals  very  low  in  the  scale,  I  shall  give 
some  additional  facts  under  Sexual  Selection,  showing  that  their 
mental  powers  are  much  higher  than  might  have  been  expected. 
The  variability  of  the  faculties  in  the  individuals  of  the  same 
species  is  an  important  point  for  us,  and  some  few  illustrations 
will  here  be  given.  But  it  would  be  superfluous  to  enter  into 
many  details  on  this  head,  for  I  have  found  on  frequent  inquiry, 
that  it  is  the  unanimous  opinion  of  all  those  who  have  long 
attended  to  animals  of  many  kinds,  including  birds,  that  the  in- 
dividuals differ  greatly  in  every  mental  characteristic.  In  what 
manner  the  mental  powers  were  first  developed  in  the  lowest 
organisms,  is  as  hopeless  an  inquiry  as  how  life  itself  first 
originated.  These  are  problems  for  the  distant  future,  if  they  are 
ever  to  be  solved  by  man. 

As  man  possesses  the  same  senses  as  the  lower  animals,  his 
fundamental  intuitions  must  be  the  same.  Man  has  also  some 
few  instincts  in  common,  as  that  of  self-preservation,  sexual  love, 
the  love  of  the  mother  for  her  new-born  offspring,  the  desire 
possessed  by  the  latter  to  suck,  and  so  forth.  But  man,  perhaps, 
has  somewhat  fewer  instincts  than  those  possessed  by  the  ani- 
mals which  come  next  to  him  in  the  series.  The  orang  in  the 
Eastern  islands,  and  the  chimpanzee  in  Africa,  build  platforms 
on  which  they  sleep;  and,  as  both  species  follow  the  same  habit, 
it  might  be  argued  that  this  was  due  to  instinct,  but  we  cannot 
feel  sure  that  it  is  not  the  result  of  both  animals  having  similar 
wants,  and  possessing  similar  powers  of  reasoning.  These  apes, 
as  we  may  assume,  avoid  the  many  poisonous  fruits  of  the  tropics, 


MENTAL    POWERS.  65' 

and  man  has  no  such  knowledge:  hut  as  our  domestic  animals, 
v/hen  taken  to  foreign  lands,  and  when  first  turned  out  in  the 
spring,  often  eat  poisonous  herbs,  which  they  afterwards  avoid, 
we  cannot  feel  sure  that  the  apes  do  not  learn  from  their  own 
experience  or  from  that  of  their  parents  what  fruits  to  select. 
It  is,  however,  certain,  as  we  shall  presently  see,  that  apes  have 
an  instinctive  dread  of  serpents,  and  probably  of  other  dangerous 
animals. 

The  fewness  and  the  comparative  simplicity  of  the  instincts  in 
the  higher  animals  are  remarkable  in  contrast  with  those  of  the 
lower  animals.  Cuvier  maintained  that  instinct  and  intelligence 
stand  in  an  inverse  ratio  to  each  other;  and  some  have  thought 
that  the  intellectual  faculties  of  the  higher  animals  have  been 
gradually  developed  from  their  instincts.  But  Pouchet,  in  an 
interesting  essay,-  has  shown  that  no  such  inverse  ratio  really 
exists.  Those  insects  which  possess  the  most  wonderful  instincts 
are  certainly  the  most  intelligent.  In  the  vertebrate  series,  the 
least  intelligent  members,  namely  fishes  and  amphibians,  do  not 
possess  complex  instincts;  and  amongst  mammals  the  animal 
most  remarkable  for  its  instincts,  namely  the  beaver,  is  highly 
intelligent,  as  will  be  admitted  by  every  one  who  has  read  Mr. 
Morgan's  excellent  work.^ 

Although  the  first  dawnings  of  intelligence,  according  to  Mr. 
Herbert  Spencer,^  have  been  developed  through  the  multiplica- 
tion and  co-ordination  of  reflex  actions,  and  although  many  of 
the  simpler  instincts  graduate  into  reflex  actions,  and  can  hardly 
be  distinguished  from  them,  as  in  the  case  of  young  animals 
sucking,  yet  the  more  complex  instincts  seem  to  have  originated 
independently  of  intelligence.  I  am,  however,  very  far  from 
wishing  to  deny  that  instinctive  actions  may  lose  their  flxe^i  and 
untaught  character,  and  be  replaced  by  others  performed  by  the 
aid  of  the  free  will.  On  the  other  hand,  some  intelligent  actions, 
after  being  performed  during  several  generations,  become  con- 
verted into  instincts  and  are  inherited,  as  when  birds  on  oceanic 
islands  learn  to  avoid  man.  These  actions  may  then  be  said 
to  be  degraded  in  character,  for  they  are  no  longer  performed 
through  reason  or  from  experience.  But  the  greater  number  of 
the  more  complex  instincts  appear  to  have  been  gained  in  a 
wholly  different  manner,  through  the  natural  selection  of  varia- 
tions of  simpler  instinctive  actions.  Such  variations  appear  to 
arise  from  the  same  unknown  causes  acting  on  the  cerebral 
organization,  which  induce  slight  variations  or  individual   dif- 

2  'L'lnstinct  chez  les  Insectes.'  'Revue  des  Deux  Mondes,'  Feb.  1870, 
p.  690. 
•^  'The  American  Beaver  and  his  Works/  1868. 
*  'The  Principles  of  Psychology,'  2nd  edit.  1870,  pp.  418-443. 
6 


66  THE  DESCENT  OP  MAN. 

ferences  in  other  parts  of  the  body;  and  these  variations,  owing 
to  our  ignorance,  are  often  said  to  arise  spontaneously.  We  can, 
I  think,  come  to  no  other  conclusion  with  respect  to  the  origin  of 
the  more  complex  instincts,  when  v/e  reflect  on  the  marvelous 
instincts  of  sterile  worker-ants  and  bees,  which  leave  no  off- 
spring to  inherit  the  effects  of  experience  and  of  modified  habits. 

Although,  as  we  learn  from  the  above-mentioned  insects  and 
the  beaver,  a  high  degree  of  intelligence  is  certainly  compatible 
with  complex  instincts,  and  although  actions,  at  first  learned 
voluntarily  can  soon  through  habit  be  performed  with  the  quick- 
ness and  certainty  of  a  reflex  action,  yet  it  is  not  improbable  that 
there  is  a  certain  amount  of  interference  between  the  devel- 
opment of  free  intelligence  and  of  instinct, — which  latter  implies 
some  inherited  modification  of  the  brain.  Little  is  known  about 
the  functions  of  the  brain,  but  we  can  perceive  that  as  the  intel- 
lectual powers  become  highly  developed,  the  various  parts  of  the 
brain  must  be  connected  by  very  intricate  channels  of  the  freest 
intercommunication;  and  as  a  consequence,  each  separate  part 
would  perhaps  tend  to  be  less  well  fitted  to  answer  to  particular 
sensations  or  associations  in  a  definite  and  inherited — that  is 
instinctive — manner.  There  seems  even  to  exist  some  relation 
between  a  low  degree  of  intelligence  and  a  strong  tendency  to  the 
formation  of  fixed,  though  not  inherited  habits;  for  as  a  sagacious 
physician  remarked  to  me,  persons  who  are  slightly  imbecile  tend 
to  act  in  everything  by  routine  or  habit;  and  they  are  rendered 
much  happier  if  this  is  encouraged. 

I  have  thought  this  digression  worth  giving,  because  we  may 
easily  underrate  the  mental  powers  of  the  higher  animals,  and 
especially  of  man,  when  we  compare  their  actions  founded  on  the 
memory  of  past  events,  on  foresight,  reason,  and  imagination, 
with  exactly  similar  actions  instinctively  performed  by  the  lower 
animals;  in  this  latter  case  the  capacity  of  performing  such 
actions  has  been  gained,  step  by  step,  through  the  variability  of 
the  mental  organs  and  natural  selection,  without  any  conscious 
intelligence  on  the  part  of  the  animal  during  each  successive 
generation.  No  doubt,  as  Mr.  Wallace  has  argued,^  much  of  the 
intelligent  work  done  by  man  is  due  to  imitation  and  not  to 
reason;  but  there  is  this  great  difference  between  his  actions 
and  many  of  those  performed  by  the  lower  animals,  namely,  that 
man  cannot,  on  his  first  trial,  mak(3,  for  instance,  a  stone  hatchet 
or  a  canoe,  through  his  power  of  imitation.  He  has  to  learn  his 
work  by  practice;  a  beaver,  on  the  other  hand,  can  make  its 
dam  or  canal,  and  a  bird  its  nest,  as  well,  or  nearly  as  well,  and 

5  'Contributions  to  the  Theory  of  Natural  Selection,'  1870,  p.  212. 


MENTAL    POWERS.  67 

a  spider  its  wonderful  web,  quite  as  well,®  the  first  time  it  tries, 
as  when  old  and  experienced. 

To  return  to  our  immediate  subject:  the  lower  animals,  like 
man,  manifestly  feel  pleasure  and  pain,  happiness  and  misery. 
Happiness  is  never  better  exhibited  than  by  young  animals,  such 
as  puppies,  kittens,  lambs,  &c.,  when  playing  together,  like  our 
own  children.  Even  insects  play  together,  as  has  been  described 
by  that  excellent  observer,  P.  Huber,'^  who  saw  ants  chasing  and 
pretending  to  bite  each  other,  like  so  many  puppies. 

The  fact  that  the  lower  animals  are  excited  by  the  same 
emotions  as  ourselves  is  so  well  established,  that  it  will  not  be 
necessary  to  weary  the  reader  by  many  details.  Terror  acts  in 
the  same  manner  on  them  as  on  us,  causing  the  muscles  to 
tremble,  the  heart  to  palpitate,  the  sphincters  to  be  relaxed,  and 
the  hair  to  stand  on  end.  Suspicion,  the  offspring  of  fear,  is 
eminently  characteristic  of  most  wild  animals.  It  is,  I  think, 
impossible  to  read  the  account  given  by  Sir  E.  Tennent,  of  the 
behavior  of  the  female  elephants,  used  as  decoys,  without  ad- 
mitting that  they  intentionally  practice  deceit,  and  well  know 
what  they  are  about.  Courage  and  timidity  are  extremely  variable 
qualities  in  the  individuals  of  the  same  species,  as  is  plainly  seen 
in  our  dogs.  Some  dogs  and  horses  are  ill-tempered,  and  easily 
turn  sulky;  others  are  good-tempered;  and  these  qualities  are  f^'^^^^ 
certainly  inherited.  Every  one  knows  how  liable  animals  are 
to  furious  rage,  and  how  plainly  they  show  it.  Many,  and  prob- 
ably true,  anecdotes  have  been  published  on  the  long-delayed  and 
artful  revenge  of  various  animals.  The  accurate  Rengger,  and 
Brehm^  state  that  the  American  and  African  monkeys  which 
they  kept  tame,  certainly  revenged  themselves.  Sir  Andrew 
Smith,  a  zoologist  whose  scrupulous  accuracy  was  known  to  many 
persons,  told  me  the  following  story  of  which  he  was  himself  an 
eye-witness;  at  the  Cape  of  Good  Hope  an  officer  had  often 
plagued  a  certain  baboon,  and  the  animal,  seeing  him  approaching 
one  Sunday  for  parade,  poured  water  into  a  hole  and  hastily 
made  some  thick  mud,  which  he  skillfully  dashed  over  the  officer 
as  he  passed  by,  to  the  amusement  of  many  bystanders.  For 
long  afterwards  the  baboon  rejoiced  and  triumphed  whenever  he 
saw  his  victim. 

The  love   of  a  dog  for  his  master  is  notorious;    as  an  old 

6  For  the  evidence  on  this  head,  see  Mr.  J.  Traherne  Moggridge's 
most  interesting  work,  'Harvesting  Ants  and  Trap-door  Spiders,'  1873, 
pp.  126,  128. 

■^  'Recherches  sur  les  Moeurs  des  Fourmis,'  1810,  p.  173. 

8  All  the  following  statements,  given  on  the  authority  of  these  two 
naturalists,  are  taken  from  Rengger's  'Naturgesch.  der  Saugethiere 
von  Paraguay,'  1830,  s.  41-57,  and  from  Brehtn's  'Thierieben,'  B.  i.  s. 


g3  THE   DESCENT  OF  MAN. 

writer  quaintly  says,^  "A  dog  is  the  only  thing  on  this  earth 
"that  luvs  you  more  than  he  luvs  himself." 

In  the  agony  of  death  a  dog  has  been  known  to  caress  his 
master,  and  every  one  has  heard  of  the  dog  suffering  under  vivi- 
section, who  licked  the  hand  of  the  operator;  this  man,  unless  the 
operation  was  fully  justified  by  an  increase  of  our  knowledge, 
or  unless  he  had  a  heart  of  stone,  must  have  felt  remorse  to  the 
last  hour  of  his  life. 

As  WhewelP"  has  well  asked,  "who  that  reads  the  touching 
"instances  of  maternal  affection,  related  so  often  of  the  women  of 
"all  nations,  and  of  the  females  of  all  animals,  can  doubt  that  the 
"principle  of  action  is  the  same  in  the  two  cases?"  We  see  mater- 
nal affection  exhibited  in  the  most  trilling  details;  thus  Rengger 
observed  an  American  monkey  (a  Cebus)  carefully  driving  away 
the  flies  which  plagued  her  infant;  and  Duvaucel  saw  a  Hylobates 
washing  the  faces  of  her  young  ones  in  a  stream.  So  intense  is 
the  grief  of  female  monkeys  for  the  loss  of  their  young,  that 
it  invariably  caused  the  death  of  certain  kinds  kept  under  confine- 
ment by  Brehm  in  N.  Africa.  Orphan  monkeys  were  always 
adopted  and  carefully  guarded  by  the  other  monlj:eys,  both  males 
and  females.  One  female  baboon  had  so  capacious  a  heart  that 
she  not  only  adopted  young  monkeys  of  other  species,  but  stole 
young  dogs  and  cats,  which  she  continually  carried  about.  Her 
kindness,  however,  did  not  go  so  far  as  to  share  her  food  with 
her  adopted  offspring,  at  which  Brehm  was  surprised,  as  his 
monkeys  always  divided  everything  quite  fairly  with  their  own 
young  ones.  An  adopted  kitten  scratched  this  affectionate  baboon, 
who  certainly  had  a  fine  intellect,  for  she  was  much  astonished  at 
being  scratched,  and  immediately  examined  the  kitten's  feet,  and 
without  more  ado  bit  off  the  claws."  In  the  Zoological  Gardens, 
I  heard  from  the  keeper  that  an  old  baboon  (C.  chacma)  had 
adopted  a  Rhesus  monkey;  but  when  a  young  drill  and  mandrill 
were  placed  in  the  cage,  she  seemed  to  perceive  that  these 
monkeys,  though  distinct  species,  were  her  nearer  relatives,  for 
she  at  once  rejected  the  Rhesus  and  adopted  both  of  them.  The 
young  Rhesus,  as  I  saw,  was  greatly  discontented  at  being  thus 
rejected,  and  it  would,  like  a  naughty  child,  annoy  and  attack  the 
young  drill  and  mandrill  whenever  it  could  do  so  with  safety;  this 
conduct  exciting  great  indignation  in  the  old  baboon.    Monkeys 

»  Quoted  by  Dr.  Lauder  Lindsay,  in  his  'Physiology  of  Mind  in  the 
Lower  Animals;'    'Journal  of  Mental  Science,'  April,  1871,  p.  38. 

10  'Bridgewater  Treatise,'  p.  263. 

"  A  critic,  without  any  grounds  ('Quarterly  Ileview>*  July,  1871,  p. 
72),  disputes  the  possibility  of  this  act  as  described  by  Brehm,  for  the 
sake  of  discrediting  my  work.  Therefore  I  tried,  and  found  that  I 
could  readily  seize  with  my  own  teeth  the  sharp  little  claws  of  a 
kitten  nearly  five  weeks  old. 


MENTAL    POWERS.  69 

will  also,  according  to  Brehm,  defend  their  master  when  attacked 
by  any  one,  as  well  as  dogs  to  whom  they  are  attached,  from  the 
attacks  of  other  dogs.  But  we  here  trench  on  the  subjects  of  sym- 
pathy and  fidelity,  to  which  I  shall  recur.  Some  of  Brehm's 
monkeys  took  much  delight  in  teasing  a  certain  old  dog  whom 
they  disliked,  as  well  as  other  animals,  in  various  ingenious  ways. 

Most  of  the  more  complex  emotions  are  common  to  the 
higher  animals  and  ourselves.  Every  one  has  seen  how  jealous 
a  dog  is  of  his  master's  affection,  if  lavished  on  any  other  crea- 
ture; and  I  have  observed  the  same  fact  with  monkeys.  This 
shows  that  animals  not  only  love,  but  have  desire  to  be  loved. 
Animals  manifestly  feel  emulation.  They  love  approbation  or 
praise;  and  a  dog  carrying  a  basket  for  his  master  exhibits  in 
a  high  degree  self-complacency  or  pride.  There  can,  I  think,  be 
no  doubt  that  a  dog  feels  shame,  as  distinct  from  fear,  and  some- 
thing very  like  modesty  when  begging  too  often  for  food.  A 
great  dog  scorns  the  snarling  of  a  little  dog,  and  this  may 
be  called  magnanimity.  Several  observers  have  stated  that 
monkeys  certainly  dislike  being  laughed  at;  and  they  sometimes 
invent  imaginary  offenses.  In  the  Zoological  Gardens  I  saw  a 
baboon  who  always  got  into  a  furious  rage  when  his  keeper  took 
out  a  letter  or  book  and  read  it  aloud  to  him;  and  his  rage  was 
so  violent  that,  as  I  witnessed  on  one  occasion,  he  bit  his  own  leg 
till  the  blood  flowed.  Dogs  show  what  may  be  fairly  called  a 
sense  of  humor,  as  distinct  from  mere  play;  if  a  bit  of  stick  or 
other  such  object  be  thrown  to  one,  he  will  often  carry  it  away 
for  a  short  distance;  and  then  squatting  down  with  it  on  the 
ground  close  before  him,  will  wait  until  his  master  comes  quite 
close  to  take  it  away.  The  dog  will  then  seize  it  and  rush  away  in 
triumph,  repeating  the  same  manoeuvre,  and  evidently  enjoying 
the  practical  joke. 

We  will  now  turn  to  the  more  intellectual  emotions  and  fac- 
ulties, which  are  very  important,  as  forming  the  basis  for  the  de- 
velopment of  the  higher  mental  powers.  Animals  manifestly 
enjoy  excitement,  and  suffer  from  ennui,  as  may  be  seen  with 
dogs,  and,  according  to  Rengger,  with  monkeys.  All  animals 
feel  Wonder,  and  many  exhibit  Curiosity.  They  sometimes  suffer 
from  this  latter  quality,  as  when  the  hunter  plays  antics  and  thus 
attracts  them;  I  witnessed  this  with  deer,  and  so  it  is  with  the 
wary  chamois,  and  with  some  kinds  of  wild-ducks.  Brehm  gives 
a  curious  account  of  the  instinctive  dread,  which  his  monkeys 
exhibited,  for  snakes;  but  their  curiosity  was  so  great  that  they 
could  not  desist  from  occasionally  satiating  their  horror  in  a 
most  human  fashion,  by  lifting  up  the  lid  of  the  box  in  which  the 
snakes  were  kept.  I  was  so  much  surprised  at  his  account,  that 
I  took  a  stuffed  and  coiled-up  snake  into  the  monkey-house  at  the 
Zoological  Gardens,  and  the  excitement  thus  caused  was  one  of 

6 


7d  THE  DESCENT  OF  MAN. 

the  most  curious  spectacles  v/^hich  I  ever  beheld.  Three  species 
of  Cercopithecus  were  the  most  alarmed;  they  dashed  about  their 
cages,  and  uttered  sharp  signal  cries  of  danger,  which  were  under- 
stood by  the  other  monkeys.  A  few  young  monkeys  and  one  old 
Anubis  baboon  alone  took  no  notice  of  the  snake.  I  then  placed 
the  stuffed  specimen  on  the  ground  in  one  of  the  larger  com- 
partments. After  a  time  all  the  monkeys  collected  round  it  in 
a  large  circle,  and  staring  intently,  presented  a  most  ludicrous  ap- 
pearance. They  became  extremely  nervous;  so  that  when  a 
wooden  ball,  with  which  they  were  familiar  as  a  plaything,  was 
accidentally  moved  in  the  straw,  under  which  it  was  partly 
hidden,  they  all  instantly  started  away.  These  monkeys  behaved 
very  differently  when  a  dead  fish,  a  mouse,^  a  living  turtle,  and 
other  new  objects  were  placed  in  their  cages;  for  though  at  first 
frightened,  they  soon  approached,  handled  and  examined  them.  I 
then  placed  a  live  snake  in  a  paper  bag,  with  the  mouth  loosely 
closed,  in  one  of  the  larger  compartments.  One  of  the  monkeys  im- 
mediately approached,  cautiously  opened  the  bag  a  little,  peeped 
in,  and  instantly  dashed  away.  Then  I  witnessed  what  Brehm  has 
described,  for  monkey  after  monkey,  with  head  raised  high  and 
turned  on  one  side,  could  not  resist  taking  a  momentary  peep 
into  the  upright  bag,  at  the  dreadful  object  lying  quietly  at  the 
bottom.  It  would  almost  appear  as  if  monkeys  had  some  notion 
of  zoological  afiinities,  for  those  kept  by  Brehm  exhibited  a 
strange,  though  mistaken,  instinctive  dread  of  innocent  lizards 
and  frogs.  An  orang,  also,  has  been  known  to  be  much  alarmed 
at  the  first  sight  of  a  turtle. ^^ 

?  The  principle  of  Imitation  is  strong  in  man,  and  especially,  as 
I  have  myself  observed,  with  savages.  In  certain  morbid  states 
"of  the  brain  this  tendency  is  exaggerated  to  an  extraordinary 
degree;  some  hemiplegic  patients  and  others,  at  the  commence- 
ment of  inflammatory  softening  of  the  brain,  unconsciously 
imitate  every  word  which  is  uttered,  whether  in  their  own  or  in 
a  foreign  language,  and  every  gesture  or  action  which  is  per- 
formed near  them."  Desor^^  has  remarked  that  no  animal  volun- 
tarily imitates  an  action  performed  by  man,  until  in  the  ascending 
scale  we  come  to  monkeys,  which  are  well  known  to  be  ridiculous 
mockers.  Animals,  however,  sometimes  imitate  each  other's 
actions:  thus  two  species  of  wolves,  which  had  been  reared  by 
dogs,  learned  to  bark,  as  does  sometimes  the  jackal,"  but  whether 

^  I  have  given  a  short  account  of  their  behavior  on  this  occasion  in 
my  'Expression  of  the  Emotions,'  p.  43. 

13  W.  C.  L.  Martin.    'Nat.  Hist,  of  Mammalia,'  1841,  p.  405. 

1*  Dr.  Bateman  'On  Apasia,'  1870,  p.  110. 

15  Quoted  by  Vogt,  'Memoire  sur  les  Microcephales,'  1867,  p.  168. 

18  'Tke  Variation  of  Animals  and  Plants  imdie'r  Domfesticatioo,'  vol.  i, 
p.  27. 


MENTAL    POWERS.  71 

this  can  be  called  voluntary  imitation  is  another  question.  Birds 
imitate  the  songs  of  their  parents,  and  sometimes  of  other  birds; 
and  parrots  are  notorious  imitators  of  any  sound  which  they  often 
hear.  Bureau  de  la  Malle  gives  an  account^'  of  a  dog  reared  by  a 
cat,  who  learned  to  imitate  the  well-known  action  of  a  cat  licking 
her  paws,  and  thus  washing  her  ears  and  face;  this  was  also 
witnessed  by  the  celebrated  naturalist  Audouin.  I  have  received 
several  confirmatory  accounts;  in  one  of  these,  a  dog  had  not  been 
suckled  by  a  cat,  but  had  been  brought  up  with  one,  together 
with  kittens,  and  had  thus  acquired  the  above  habit,  which  he 
ever  afterwards  practiced  during  his  life  of  thirteen  years.  Bureau 
de  la  Malle's  dog  likewise  learned  from  the  kittens  to  play  with  a 
ball  by  rolling  it  about  with  his  fore  paws,  and  springing  on  it. 
A  correspondent  assures  me  that  a  cat  in  his  house  used  to  put 
her  paws  into  jugs  of  milk  having  too  narrow  a  mouth  for  her 
head.  A  kitten  of  this  cat  soon  learned  the  same  trick,  and  prac- 
ticed it  ever  afterwards,  whenever  there  was  an  opportunity. 

The  parents  of  many  animals,  trusting  to  the  principle  of 
imitation  in  their  young,  and  more  especially  to  their  instincdve 
or  inherited  tendencies,  may  be  said  to  educate  them.  We  see 
this  when  a  cat  brings  a  live  mouse  to  her  kittens;  and  Bureau 
de  la  Malle  has  given  a  curious  account  (in  the  paper  above 
quoted)  of  his  observations  on  hawks  which  taught  their  young 
dexterity,  as  well  as  judgment  of  distances,  by  first  dropping 
through  the  air  dead  mice  and  sparrows,  which  the  young  gen- 
erally failed  to  catch,  and  then  bringing  them  live  birds  and 
letting  them  loose. 

Hardly  any  faculty  is  more  important  for  the  intellectual 
progress  of  man  than  Attention.  Animals  clearly  manifest  this 
power,  as  when  a  cat  watches  by  a  hole  and  prepares  to  spring 
on  its  prey.  Wild  animals  sometimes  become  so  absorbed  when 
thus  engaged,  that  they  may  be  easily  approached.  Mr.  Bartlett 
has  given  me  a  curious  proof  how  variable  this  faculty  is  in 
monkeys.  A  man  who  trains  monkeys  to  act  in  plays,  used  to 
purchase  common  kinds  from  the  Zoological  Society  at  the  price 
of  five  pounds  for  each;  but  he  offered  to  give  double  the  price, 
if  he  might  keep  three  or  four  of  them  for  a  few  days,  in  order 
to  select  one.  When  asked  how  he  could  possibly  learn  so  soon, 
whether  a  particular  monkey  would  turn  out  a  good  actor,  he 
answered  that  it  all  depended  on  their  power  of  attention.  If, 
when  he  was  talking  and  explaining  anything  to  a  monkey,  its 
attention  was  easily  distracted,  as  by  a  fly  on  the  wall  or  other 
trifling  object,  the  case  was  hopeless.  If  he  tried  by  punishment 
to  make  an  inattentive  monkey  act,  it  turned  sulky.     On  the 


I''  'Annales  des  Sc.  Nat.'  (Ist  Series),  torn.  xaai.  p.  39-7. 


72  THE   DESCENT  OF  MAN. 

other  hand,  a  monkey  which  carefully  attended  to  him  could 
always  be  trained. 

It  is  almost  superfluous  to  state  that  animals  have  excellent 
Memories  for  persons  and  places.  A  baboon  at  the  Cape  of  Good 
Hope,  as  I  have  been  informed  by  Sir  Andrew  Smith,  recognized 
him  with  joy  after  an  absence  of  nine  months.  I  had  a  dog  who 
was  savage  and  averse  to  all  strangers,  and  I  purposely  tried  his 
memory  after  an  absence  of  five  years  and  two  days.  I  went 
near  the  stable  where  he  lived,  and  shouted  to  him  in  my  old 
manner;  he  showed  no  joy,  but  instantly  followed  me  out  walk- 
ing, and  obeyed  me,  exactly  as  if  I  had  parted  with  him  only 
half  an  hour  before.  A  train  of  old  associations,  dormant  during 
five  years,  had  thus  been  instantaneously  awakened  in  his  mind. 
Even  ants,  as  P.  Huber^*  has  clearly  shown,  recognized  their 
fellow-ants  belonging  to  the  same  community  after  a  separation 
of  four  months.  Animals  can  certainly  by  some  means  judge  of 
the  intervals  of  time  between  recurrent  events. 
(_.  The  Imagination  is  one  of  the  highest  prerogatives  of  man. 
By  this  faculty  he  unites  former  images  and  ideas,  independently 
of  the  will,  and  thus  creates  brilliant  and  novel  results.  A  poet, 
as  Jean  Paul  Richter  remarks,^^  "who  must  reflect  whether  he 
"shall  make  a  character  say  yes  or  no — to  the  devil  with  him; 
"he  is  only  a  stupid  corpse."  Dreaming  gives  us  the  best  notion 
of  this  power;  as  Jean  Paul  again  says,  "The  dream  is  an  in- 
"voluntary  art  of  poetry."  The  value  of  the  products  of  our 
imagination  depends  of  course  on  the  number,  accuracy,  and 
clearness  of  our  impressions,  on  our  judgment  and  taste  in  select- 
ing or  rejecting  the  involuntary  combinations,  and  to  a  certain 
extent  on  our  power  of  voluntarily  combining  them.  As  dogs, 
cats,  horses,  and  probably  all  the  higher  animals,  even  birds-" 
have  vivid  dreams,  and  this  is  shown  by  their  movements  and 
the  sounds  uttered,  we  must  admit  that  they  possess  some  power 
of  imagination.  There  must  be  something  special,  which  causes 
dogs  to  howl  in  the  night,  and  especially  during  moonlight,  in 
that  remarkable  and  melancholy  manner  called  baying.  All  dogs 
do  not  do  so;  and,  according  to  Houzeau,-^  they  do  not  then 
look  at  the  moon,  but  at  some  fixed  point  near  the  horizon. 
Houzeau  thinks  that  their  imaginations  are  disturbed  by  the 
vague  outlines  of  the  surrounding  objects,  and  conjure  up  before 

18  'Les  Mceurs  des  Fourmis,'  1810,  p.  150. 

1^  Quoted  in  Dr.  Maudsley's  Physiology  and  Pathology  of  Mind,'  1868, 
pp.  19,  220. 

20  Dr.  Jerdon,  'Birds  of  India,'  vol.  i.  1862,  p.  xxi.  Houzeau  says  that 
his  parokeets  and  canary-birds  dreamt:  'Facultes  Mental es,'  torn.  ii. 
p.  136. 

21  'Facultes  Mentales  des  Anknaux,'  1872,  torn.  ii.  p.  181. 


MENTAL    POWERS.  73 

them  fantastic  images:    if  this  be  so,  their  feelings  may  almost 
be  called  superstitious. 

Of  all  the  faculties  of  the  human  mind,  it  will,  I  presume,  be 
admitted  that  Reason  stands  at  the  summit.  Only  a  few  persons 
now  dispute  that  animals  possess  some  power  of  reasoning. 
Animals  may  constantly  be  seen  to  pause,  deliberate,  and  resolve. 
It  is  a  significant  fact,  that  the  more  the  habits  of  any  particular 
animal  are  studied  by  a  naturalist,  the  more  he  attributes  to 
reason  and  the  less  to  unlearned  instincts.--  In  future  chapters 
we  shall  see  that  some  animals  extremely  low  in  the  scale  appar- 
ently display  a  certain  amount  of  reason.  No  doubt  it  is  often 
difficult  to  distinguish  between  the  power  of  reason  and  that  of 
instinct.  For  instance.  Dr.  Hayes,  in  his  work  on  'The  Open 
Polar  Sea,'  repeatedly  remarks  that  his  dogs,  instead  of  continu- 
ing to  draw  the  sledges  in  a  compact  body,  diverged  and  separ- 
ated when  they  came  to  thin  ice,  so  that  their  weight  might  be 
more  evenly  distributed.  This  was  often  the  first  warning 
which  the  travelers  received  that  the  ice  was  becoming  thin  and 
dangerous.  Now,  did  the  dogs  act  thus  from  the  experience  of 
each  individual,  or  from  the  example  of  the  older  and  wiser  dogs, 
or  from  an  inherited  habit,  that  is  from  instinct?  This  instinct, 
may  possibly  have  arisen  since  the  time,  long  ago,  when  dogs 
were  first  employed  by  the  natives  in  drawing  their  sledges;  or 
the  Arctic  wolves,  the  parent-stock  of  the  Esquimaux  dog,  may 
have  acquired  an  instinct,  impelling  them  not  to  attack  their 
prey  in  a  close  pack,  when  on  thin  ice. 

We  can  only  judge  by  the  circumstances  under  which  actions 
are  performed,  whether  they  are  due  to  instinct,  or  to  reason,  or 
to  the  mere  association  of  ideas:  this  latter  principle,  however, 
is  intimately  connected  with  reason.  A  curious  case  has  been 
given  by  Prof.  Mobius,-^  of  a  pike,  separated  by  a  plate  of  glass 
from  an  adjoining  aquarium  stocked  with  fish,  and  who  often 
dashed  himself  with  such  violence  against  the  glass  in  trying  to 
catch  the  other  fishes,  that  he  was  sometimes  completely 
stunned.  The  pike  went  on  thus  for  three  months,  but  at  last 
learned  caution,  and  ceased  to  do  so.  The  plate  of  glass  was  then 
removed,  but  the  pike  would  not  attack  these  particular  fishes, 
though  he  would  devour  others  which  were  afterward  intro- 
duced; so  strongly  was  the  idea  of  a  violent  shock  associated 
in  his  feeble  mind  with  the  attempt  on  his  former  neighbors. 
If  a  savage,  who  had  never  seen  a  large  plate-glass  window, 
were  to  dash  himself  even  once  against  it,  he  would  for  a  long 

22  Mr.  Li.  H.  Morgan's  work  on  'The  American  Beaver,'  1868,  offers 
a  good  illustration  of  this  remark.  I  cannot  help  thinking,  however, 
that  he  goes  too  far  in  underrating  the  power  of  instinct. 

-^  'Die  Bewegungen  der  Thiere,'  &c.,  1873,  p.  11. 


74  THE  DESCENT  OF  MAN. 

time  afterwards  associate  a  shock  with  a  window-frame;  but 
very  differently  from  the  pike,  he  would  probably  reflect  on  the 
nature  of  the  impediment,  and  be  cautious  under  analogous  cir- 
cumstances. Now  with  monkeys,  as  we  shall  presently  see,  a 
painful  or  merely  a  disagreeable  impression,  from  an  action  once 
performed,  is  sometimes  sufficient  to  prevent  the  animal  from 
repeating  it.  If  we  attribute  this  difference  between  the  monkey 
and  the  pike  solely  to  the  association  of  ideas  being  so  much 
stronger  and  more  persistent  in  the  one  than  the  other,  though 
the  pike  often  received  much  the  more  severe  injury,  can  we 
maintain  in  the  case  of  man  that  a  similar  difference  implies  the 
possession  of  a  fundamentally  different  mind? 

Houzeau  relates-*  that,  whilst  crossing  a  wide  and  arid  plain 
in  Texas,  his  two  dogs  suffered  greatly  from  thirst,  and  that 
between  thirty  and  forty  times  they  rushed  down  the  hollows 
to  search  for  water.  These  hollows  were  not  valleys,  and  there 
were  no  trees  in  them,  or  any  other  difference  in  the  vegetation, 
and  as  they  were  absolutely  dry  there  could  have  been  no 
smell  of  damp  earth.  The  dogs  behaved  as  if  they  knew  that 
a  dip  in  the  ground  offered  them  the  best  chance  of  finding 
water,  and  Houzeau  has  often  witnessed  the  same  behavior  in 
other  animals. 

I  have  seen,  as  I  daresay  have  others,  that  when  a  small 
object  is  thrown  on  the  ground  beyond  the  reach  of  one  of  the 
elephants  in  the  Zoological  Gardens,  he  blows  through  his  trunk 
on  the  ground  beyond  the  object,  so  that  the  current  reflected 
on  all  sides  may  drive  the  object  within  his  reach.  Again  a  well- 
known  ethnologist,  Mr.  Westropp,  informs  me  that  he  observed  in 
Vienna  a  bear  deliberately  making  with  his  paw  a  current  in  some 
water,  which  was  close  to  the  bars  of  his  cage,  so  as  to  draw 
a  piece  of  floating  bread  within  his  reach.  These  actions  of  the 
elephant  and  bear  can  hardly  be  attributed  to  instinct  or  inherited 
habit,  as  they  would  be  of  little  use  to  an  animal  in  a  state  of 
nature.  Now,  what  is  the  difference  between  such  actions,  when 
performed  by  an  uncultivated  man,  and  by  one  of  the  higher 
animals? 

The  savage  and  the  dog  have  often  found  water  at  a  low  level, 
and  the  coincidence  under  such  circumstances  has  become  asso- 
ciated in  their  minds.  A  cultivated  man  would  perhaps  make 
some  general  proposition  on  the  subject;  but  from  all  that  we 
know  of  savages  it  is  extremely  doubtful  whether  they  would  do 
so,  and  a  dog  certainly  would  not.  But  a  savage,  as  well  as  a 
dog,  would  search  in  the  same  way,  though  frequently  dis- 
appointed; and  in  both  it  seems  to  be  equally  an  act  of  reason, 
whether  or  not  any  general  proposition  on  the  subject  is  con- 

2*  'Facultes  Mental es  des  Animaux,'  1872,  torn.  ii.  p.  265. 


MENTAL    POWERS.  75 

sciously  placed  before  the  mind.-''  The  same  would  apply  to 
the  elephant  and  the  bear  making  currents  in  the  air  or  water. 
Tlie  savage  would  certainly  neither  know  nor  care  by  what  law, 
the  desired  movements  were  effected;  yet  his  act  would  be 
guided  by  a  rude  process  of  reasoning,  as  surely  as  would  a 
philosopher  in  his  longest  chain  of  deductions.  There  would  no 
doubt  be  this  difference  between  him  and  one  of  the  higher 
animals',  that  he  would  take  notice  of  much  slighter  circum- 
stances and  conditions,  and  would  observe  any  connection  be- 
tween them  after  much  less  experience,  and  this  would  be  of 
paramount  importance.  I  kept  a  daily  record  of  the  actions  of 
one  of  my  infants,  and  when  he  was  about  eleven  months  old, 
and  before  he  could  speak  a  single  word,  I  was  continually 
struck  with  the  greater  quickness,  with  which  all  sorts  of  objects 
and  sounds  were  associated  together  in  his  mind,  compared  with 
that  of  the  most  intelligent  dogs  I  ever  knew.  But  the  higher 
animals  differ  in  exactly  the  same  way  in  this  power  of  associa- 
tion from  those  low  in  the  scale,  such  as  the  pike,  as  well  as  in 
that  of  drawing  inferences  and  of  observation. 

The  promptings  of  reason,  after  very  short  experience,  are  well 
shown  by  the  following  actions  of  American  monkeys,  which 
stand  low  in  their  order.  Rengger,  a  most  careful  observer, 
states  that  when  he  first  gave  eggs  to  his  monkeys  in  Paraguay, 
they  smashed  them,  and  thus  lost  much  of  their  contents;  after- 
wards they  gently  hit  one  end  against  some  hard  body,  and 
picked  off  the  bits  of  shell  with  their  fingers.  After  cutting 
themselves  only  once  with  any  sharp  tool  they  would  not  touch 
it  again,  or  would  handle  it  with  the  greatest  caution.  Lumps 
of  sugar  were  often  given  them  wrapped  up  in  paper;  and 
Rengger  sometimes  put  a  live  wasp  in  the  paper,  so  that  in 
hastily  unfolding  it  they  got  stung;  after  this  had  once  hap- 
pened, they  always  first  held  the  packet  to  their  ears  to  detect 
any  movement  within.-*^ 

The  following  cases  relate  to  dogs.  Mr.  Colquhoun-'  winged 
two  wild-ducks,  which  fell  on  the  further  side  of  a  stream;  his 
retriever  tried  to  bring  over  both  at  once,  but  could  not  succeed; 

25  Prof.  Huxley  has  analyzed  with  admirable  clearness  the  mental 
steps  by  which  a  man,  as  well  as  a  dog-,  arrives  at  a  conclusion  in  a 
case  analogous  to  that  given  in  my  text.  See  his  article,  'Mr.  Darwin's 
Critics,'  in  the  'Contemporary  Review,'  Nov.  1871,  p.  462,  and  in  his 
'Critiques  and  Essays,'  1873,  p.  279. 

^  Mr.  Belt,  in  his  most  interesting  work,  'The  Naturalist  in  Nicara^ 
gua,'  1874  (p.  119),  likewise  describes  various  actions  of  a  tamed  Cebus, 
which,  I  think,  clearly  show  that  this  animal  possessed  some  reasoning 
power. 

-'  'The  Moor  and  the  Loch,'  p.  45.  Col.  Hutchinson  on  'Dog  Break- 
ing,' 1850,  p.  46. 


76  THE   DESCENT  OF  MAN. 

she  then,  though  never  before  known  to  ruffle  a  feather, 
deliberately  killed  one,  brought  over  the  other,  and  returned 
for  the  dead  bird.  Col.  Hutchinson  relates  that  two  partridges 
were  shot  at  once,  one  being  killed,  the  other  wounded;  the 
latter  ran  away,  and  was  caught  by  the  retriever,  who  on  her 
return  came  across  the  dead  bird;  "she  stopped,  evidently 
"greatly  puzzled,  and  after  one  or  two  trials,  finding  she  could 
"nqt  take  it  up  without  permitting  the  escape  of  the  winged 
"bird,  she  considered  a  moment,  then  deliberately  murdered  it 
"by  giving  it  a  severe  crunch,  and  afterwards  brought  away 
"both  together.  This  was  the  only  known  instance  of  her 
"ever  having  willfully  injured  any  game."  Here  we  have  reason 
though  not  quite  perfect,  for  the  retriever  might  have  brought 
the  wounded  bird  first  and  then  returned  for  the  dead  one,  as  in 
the  case  of  the  two  wild-ducks.  I  give  the  above  cases,  as  resting 
on  the  evidence  of  two  independent  witnesses,  and  because  in  both 
instances  the  retrievers,  after  deliberation,  broke  through  a  habit 
which  is  inherited  by  them  (that  of  not  killing  the  game  re- 
trieved), and  because  they  show  how  strong  their  reasoning  fac- 
ulty must  have  been  to  overcome  a  fixed  habit. 

I  will  conclude  by  quoting  a  remark  by  the  illustrious  Hum- 
boldt.-^ "The  muleteers  in  S.  America  say,  *I  will  not  give 
"  'you  the  mule  whose  step  is  easiest,  but  la  mas  racional, — the 
"  'one  that  reasons  best;'  "  and  as  he  adds,  "this  popular  expres- 
"sion,  dictated  by  long  experience,  combats  the  system  of  ani- 
"mated  machines,  better  perhaps  than  all  the  arguments  of  specu- 
"lative  philosophy."  Nevertheless  some  writers  even  yet  deny 
that  the  higher  animals  possess  a  trace  of  reason;  and  they  en- 
deavor to  explain  away,  by  what  appears  to  be  mere  verbiage,^ 
all  such  facts  as  those  above  given. 

It  has,  I  think,  now  been  shown  that  man  and  the  higher  ani- 
mals, especially  the  Primates,  have  some  few  instincts  in  common. 
All  have  the  same  senses,  intuitions,  and  sensations, — similar 
passions,  affections,  and  emotions,  even  the  more  complex  ones, 
such  as  jealousy,  suspicion,  emulation,  gratitude,  and  magnanim- 

28  'Personal  Narrative,'  Eng-.  translat.  vol.  iii.  p.  106. 

^  I  am  glad  to  find  that  so  acute  a  reasoner  as  Mr.  Leslie  Stephen 
('Darwinism  and  Divinity,  Essays  on  Free- thinking,'  1873,  p.  SO),  in 
speaking  of  the  supposed  impassable  harrier  between  the  minds  of  man 
and  the  lower  animals,  says,  "The  distinctions,  indeed,  which  have 
"been  drawn,  seem  to  us  to  rest  upon  no  better  foundation  than  a 
"great  many  other  metaphysical  distinctions;  that  is  the  assumption 
"that  because  you  can  give  two  things  different  names,  they  must 
"therefore  have  different  natures.  It  is  difficult  to  understand  how 
"anybody  who  has  ever  kept  a  dog,  or  seen  an  elephant,  can  have  any 
"doubts  as  to  an  animal's  power  of  performing  the  essential  processes 
*'of  reasoning." 


MENTAL    POWERS.  77 

ity;  they  practice  deceit  and  are  revengeful;  they  are  sometimes 
susceptible  to  ridicule,  and  even  have  a  sense  of  humor;  they 
teel  wonder  and  curiosity;  they  possess  the  same  faculties  of 
imitation,  attention,  deliberation,  choice,  memory,  imagination, 
the  association  of  ideas,  and  reason,  though  in  very  different  de- 
grees. The  individuals  of  the  same  species  graduate  in  intellect 
from  absolute  imbecility  to  high  excellence.  They  are  also 
liable  to  insanity,  though  far  less  often  than  in  the  case  of  man.^" 
Nevertheless,  many  authors  have  insisted  that  man  is  divided  by 
an  insuperable  barrier  from  all  the  lov/er  animals  in  his  mental 
faculties.  I  formerly  made  a  collection  of  above  a  score  of  such 
aphorisms,  but  they  are  almost  worthless,  as  their  wide  difference 
and  number  prove  the  difficulty,  if  not  the  impossibility,  of  the 
attempt.  It  has  been  asserted  that  man  alone  is  capable  of 
progressive  improvement;  that  he  alone  makes  use  of  tools  or 
fire,  domesticates  other  animals,  or  possesses  property;  that  no 
animal  has  the  power  of  abstraction,  or  of  forming  general  con- 
cepts, is  self-conscious  and  comprehends  itself;  that  no  animal 
employs  language;  that  man  alone  has  a  sense  of  beauty,  is  liable 
to  caprice,  has  the  feeling  of  gratitude,  mystery,  &c.;  believes  in 
God,  or  is  endowed  with  a  conscience.  I  will  hazard  a  few  re- 
marks on  the  more  important  and  interesting  of  these  points. 

Archbishop  Sumner  formerly  maintained^^  that  man  alone  is 
capable  of  progressive  improvement.  That  he  is  capable  of  in- 
comparably greater  and  more  rapid  improvement  than  is  any 
other  animal,  admits  of  no  dispute;  and  this  is  mainly  due  to 
his  power  of  speaking  and  handing  down  his  acquired  knowledge. 
With  animals,  looking  first  to  the  individual,  every  one  who  has 
had  any  experience  in  setting  traps,  knows  that  young  animals 
can  be  caught  much  more  easily  than  old  ones;  and  they 
can  be  much  more  easily  approached  by  an  enemy.  Even 
with  respect  to  old  animals,  it  is  impossible  to  catch  many 
in  the  same  place  and  in  the  same  kind  of  trap,  or  to 
destroy  them  by  the  same  kind  of  poison;  yet  it  is  im- 
probable that  all  should  have  partaken  of  the  poison,  and 
impossible  that  all  should  have  been  caught  in  a  trap.  They 
must  learn  caution  by  seeing  their  brethren  caught  or  poi- 
soned. In  North  America,  where  the  fur-bearing  animals  have 
long  been  pursued,  they  exhibit  according  to  the  unanimous 
testimony  of  all  observers,  an  almost  incredible  amount  of 
sagacity,  caution  and  cunning;  but  trapping  has  been  there  so 
long  carried  on,  that  inheritance  may  possibly  have  come  into 
play.    I  have  received  several  accounts  that  when  telegraphs  are 

^°  See  'Madness  in  Animals,'  by  Dr.  W.  Lauder  Lindsay,  in  'Journal 
of  Mental  Science,'  July  1871. 
31  Quoted  by  Sir  C.  Lyell,  'Antiquity  of  Man,'  p.  497. 


78  THE  DESCENT  OF  MAN. 

first  set  up  in  any  district,  many  birds  kill  themselves  by  flying 
against  the  v/ires,  but  that  in  the  course  of  a  very  few  years  they 
learn  to  avoid  this  danger,  by  seeing,  as  it  would  appear,  their 
comrades  killed."- 

If  we  look  to  successive  generations,  or  to  the  race,  there  is  no 
doubt  that  birds  and  other  animals  gradually  both  acquire  and 
lose  caution  in  relation  to  man  or  other  enemies  ;^^  and  this 
caution  is  certainly  in  chief  part  an  inherited  habit  or  instinct, 
but  in  part  the  result  of  individual  experience.  A  good  observer, 
Leroy,^*  states,  that  in  districts  where  foxes  are  much  hunted, 
the  young,  on  first  leaving  their  burrows,  are  incontestably  much 
more  wary  than  the  old  ones  in  districts  where  they  are  not  much 
disturbed. 

Our  domestic  dogs  are  descended  from  wolves  and  jackals,-^ 
and  though  they  may  not  have  gained  in  cunning,  and  may  have 
lost  in  wariness  and  suspicion,  yet  they  have  progressed  in 
certain  moral  qualities,  such  as  in  affection,  trust-worthiness, 
temper,  and  probably  in  general  intelligence.  The  common  rat  has 
conquered  and  beaten  several  other  species  throughout  Europe, 
in  parts  of  North  America,  New  Zealand,  and  recently  in  Formosa, 
as  well  as  on  the  mainland  of  China.  Mr.  Swinhoe,^"  who  de- 
scribes these  two  latter  cases,  attributes  the  victory  of  the  com- 
mon rat  over  the  large  Mus  cominga  to  its  superior  cunning;  and 
this  latter  quality  may  probably  be  attributed  to  the  habitual 
exercise  of  all  its  faculties  in  avoiding  extirpation  by  man,  as 
well  as  to  nearly  all  the  less  cunning  or  weak-minded  rats  having 
been  continuously  destroyed  by  him.  It  is,  however,  possible  that 
the  success  of  the  common  rat  may  be  due  to  its  having  pos- 
sessed greater  cunning  than  its  fellow-species,  before  it  became 
associated  with  man.  To  maintain,  independently  of  any  direct 
evidence,  that  no  animal  during  the  course  of  ages  has  progressed 
in  intellect  or  other  mental  faculties,  is  to  beg  the  question  of 
the  evolution  of  species.  We  have  seen  that,  according  to  Lartet, 
existing  mammals  belonging  to  several  orders  have  larger  brains 
than  their  ancient  tertiary  prototypes. 

It  has  often  been  said  that  no  animal  uses  any  tool;  but 
the  chimpanzee  in  a  state  of  nature  cracks  a  native  fruit,  some- 

32  For  additional  evidence,  with  details,  see  M.  Houzeau,  'Les  Facultes 
Mentales,'  torn.  ii.  1872,  p.  147. 

S3  See,  with  respect  to  birds  on  oceanic  islands,  my  'Journal  of  Re- 
searches during  the  voyage  of  the  "Beagle"  '  1845,  p.  398.  'Origin  of 
Species,'  5th  edit.  p.  260. 

34  'Lettres  Phil,  sur  I'lntelligence  des  Animaux,'  nouvelle  edit.  1892, 
p.  86. 

33  See  the  evidence  on  this  head  in  chap.  i.  vol.  i.  'On  the  Variation 
of  Animals  and  Plants  under  Domestication.' 

38  'Proc.  Zoolog.  Soc'  1864,  p.  186. 


MENTAL    POWERS.  79 

what  like  a  walnut,  with  a  stone.^^  Rengger^*  easily  taught  an 
American  monkey  thus  to  break  open  hard  palm-nuts;  and  after- 
wards of  its  own  accord,  it  used  stones  to  open  other  kinds  of 
nuts,  as  well  as  boxes.  It  thus  also  removed  the  soft  rind  of  fruit 
that  had  a  disagreeable  flavor.  Another  monkey  was  taught  to 
open  the  lid  of  a  large  box  with  a  stick,  and  afterwards  it  used 
the  stick  as  a  lever  to  move  heavy  bodies;  and  I  have  myself  seen 
a  young  orang  put  a  stick  into  a  crevice,  slip  his  hand  to  the  other 
end,  and  use  it  in  the  proper  manner  as  a  lever.  The  tamed  ele- 
phants in  India  are  well  known  to  break  off  branches  of  trees 
and  use  them  to  drive  away  the  flies;  and  this  same  act  has  been 
observed  in  an  elephant  in  a  state  of  nature.^^"  I  have  seen  a  young 
orang,  when  she  thought  she  was  going  to  be  whipped,  cover  and 
protect  herself  with  a  blanket  or  straw.  In  these  several  cases 
stones  and  sticks  were  employed  as  implements;  but  they  are 
likewise  used  as  weapons.  Brehm^  states,  on  the  authority  of  the 
well-known  traveler  Schimper,  that  in  Abyssinia  when  the 
baboons  belonging  to  one  species  (C.  gelada)  descend  in  troops 
from  the  mountains  to  plunder  the  fields,  they  sometimes  en- 
counter troops  of  another  species  (C.  hamadryas),  and  then  a 
fight  ensues.  The  Geladas  roll  down  great  stones,  which  the  Ham- 
adryas try  to  avoid,  and  then  both  species,  making  a  great  up- 
roar, rush  furiously  against  each  other.  Brehm,  when,  accom- 
panying the  Duke  of  Coburg-Gotha,  aided  in  an  attack  with  fire- 
arms on  a  troop  of  baboons  in  the  pass  of  Mensa  in  Abyssinia. 
The  baboons  in  return  rolled  so  many  stones  down  the  mountain, 
some  as  large  as  a  man's  head,  that  the  attackers  had  to  beat  a 
hasty  retreat;  and  the  pass  was  actually  closed  for  a  time  against 
the  caravan.  It  deserves  notice  that  these  baboons  thus  acted 
in  concert.  Mr.  Wallace*^  on  three  occasions  saw  female  orangs, 
accompanied  by  their  young,  "breaking  off  branches  and  the 
"great  spiny  fruit  of  the  Durian  tree,  with  every  appearance  of 
"rage;  causing  such  a  shower  of  missiles  as  effectually  kept  us 
"from  approaching  too  near  the  tree."  As  I  have  repeatedly  seen, 
a  chimpanzee  will  throw  any  object  at  hand  at  a  person  who 
offends  him;  and  the  before  mentioned  baboon  at  the  Cape  of 
Good  Hope  prepared  mud  for  the  purpose. 

In  the  Zoological  Gardens,  a  monkey,  which  had  weak  teeth, 
used  to  break  open  nuts  with  a  stone;  and  I  was  assured  by  the 
keepers  that  after  using  the  stone,  he  hid  it  in  the  straw,  and 

37  Savag-e  and  Wyman  in  'Boston  Journal  of  Nat.  Hist.'  vol.  iv.  1843- 
44,  p.  383. 
28  'Saug-ethiere  von  Paraguay,'  1830,  s.  51-56. 
s9  The  'Indian  Field,'  March  4,  1871. 
«  'Thierleben,'  B.  i.  s.  79,  82. 
*i  'The  Malay  Archipelag-o,*  vol.  i.  1869,  p.  87. 


go  THE  DESCENT  OF  MAN. 

would  not  let  any  other  monkey  touch  it.  Here,  then,  we  have 
the  idea  of  property;  but  this  idea  is  common  to  every  dog  with 
a  bone,  and  to  most  or  all  birds  with  their  nests. 

The  Duke  of  Argyll'^  remarks,  that  the  fashioning  of  an  imple- 
ment for  a  special  purpose  is  absolutely  peculiar  to  man;  and  he 
considers  that  this  forms  an  immeasurable  gulf  between  him  and 
the  brutes.  This  is  no  doubt  a  very  important  distinction;  but 
there  appears  to  me  much  truth  in  Sir  J.  Lubbock's  suggestion,"^ 
that  when  primeval  man  first  used  flint-stones  for  any  purpose, 
he  would  have  accidentally  splintered  them,  and  would  then  have 
used  the  sharp  fragments.  From  this  step  it  would  be  a  small 
one  to  break  the  flints  on  purpose,  and  not  a  very  wide  step  to 
fashion  them  rudely.  This  latter  advance,  however,  may  have 
taken  long  ages,  if  we  may  judge  by  the  immense  interval  of 
time  which  elapsed  before  the  men  of  the  neolithic  period  took  to 
grinding  and  polishing  their  stone  tools.  In  breaking  the  flints, 
as  Sir  J.  Lubbock  likewise  remarks,  sparks  would  have  been 
emitted,  and  in  grinding  them  heat  would  have  been  evolved: 
thus  the  two  usual  methods  of  "obtaining  fire  may  have  orig- 
"inated."  The  nature  of  fire  would  have  been  known  in  the 
many  volcanic  regions  where  lava  occasionally  flows  through  for- 
ests. The  anthropomorphous  apes,  guided  probably  by  instinct, 
build  for  themselves  temporary  platforms;  but  as  many  instincts 
are  largely  controlled  by  reason,  the  simpler  ones,  such  as  this 
of  building  a  platform,  might  readily  pass  into  a  voluntary  and 
conscious  act.  The  orang  is  known  to  cover  itself  at  night  with 
the  leaves  of  the  Pandanus;  and  Brehm  states  that  one  of  his 
baboons  used  to  protect  itself  from  the  heat  of  the  sun  by  throw- 
ing a  straw-mat  over  its  head.  In  these  several  habits,  we  prob- 
ably see  the  first  steps  towards  some  of  the  simpler  arts,  such 
as  rude  architecture  and  dress,  as  they  arose  amongst  the  early 
progenitors  of  man. 

Abstraction^  Oeneral  Co7iceptions,  Self -consciousness,  Mental 
Individuality.  — It  would  be  very  difiicult  for  any  one  with  even 
much  more  knowledge  than  I  possess,  to  determine  how  far  ani- 
mals exhibit  any  traces  of  these  high  mental  powers.  This  diffi- 
culty arises  from  the  impossibility  of  judging  what  passes  through 
the  mind  of  an  animal;  and  again,  the  fact  that  writers  differ  to 
a  great  extent  in  the  meaning  which  they  attribute  to  the  above 
terms,  causes  a  further  difficulty.  If  one  may  judge  from  various 
articles  which  have  been  published  lately,  the  greatest  stress 
seems  to  be  laid  on  the  supposed  entire  absence  in  animals  of  the 
power  of  abstraction,  or  of  forming  general  concepts.    But  when 


*2  'Primeval  Man,'  1869,  pp.  145,  147. 
*s  'Prehistoric  Times,'  1865,  p.  473,  &c. 


MENTAL    POWERS.  81 

a  dog  sees  another  dog  at  a  distance,  it  is  often  clear  that  he  per- 
ceives that  it  is  a  dog  in  the  abstract;  for  when  he  gets  nearer  his 
whole  manner  suddenly  changes,  if  the  other  dog  be  a  friend. 
A  recent  writer  remarks,  that  in  all  such  cases  it  is  a  pure  as- 
sumption to  assert  that  the  mental  act  is  not  essentially  of  the 
same  nature  in  the  animal  as  in  man.  If  either  refers  what  he 
perceives  with  his  senses  to  a  mental  concept,  then  so  do  both.** 
When  I  say  to  my  terrier,  in  an  eager  voice  (and  I  have  made  the 
trial  many  times),  "Hi,  hi,  where  is  it?"  she  at  once  takes  it  as 
a  sign  that  something  is  to  be  hunted,  and  generally  first  looks 
quickly  all  around,  and  then  rushes  into  the  nearest  thicket,  to 
scent  for  any  game,  but  finding  nothing,  she  looks  up  into  any 
neighboring  tree  for  a  squirrel.  Now  do  not  these  actions  clearly 
show  that  she  had  in  her  mind  a  general  idea  or  concept  that  some 
animal  is  to  be  discovered  and  hunted? 

It  may  be  freely  admitted  that  no  animal  is  self-conscious,  if 
by  this  term  it  is  implied,  that  he  refiects  on  such  points,  as 
whence  he  comes  or  whither  he  will  go,  or  v/hat  is  life  and  death, 
and  so  forth.  But  how  can  we  feel  sure  that  an  old  dog  with  an 
excellent  memory  and  some  power  of  imagination,  as  shown  by 
his  dreams,  never  reflects  on  his  past  pleasures  or  pains  in  the 
chase?  And  this  would  be  a  form  of  self-consciousness.  On  the 
other  hand,  as  Biichner*^  has  remarked,  how  little  can  the  hard- 
worked  wife  of  a  degraded  Australian  savage,  who  uses  very 
few  abstract  words,  and  cannot  count  above  four,  exert  her  self- 
consciousness,  or  reflect  on  the  nature  of  her  own  existence.  It 
is  generally  admitted,  that  the  higher  animals  possess  memory, 
attention,  association,  and  even  some  imagination  and  reason. 
If  these  powers,  which  differ  much  in  different  animals,  are 
capable  of  improvement,  there  seems  no  great  improbability  in 
more  complex  faculties,  such  as  the  higher  forms  of  abstraction, 
and  self-consciousness,  &c.,  having  been  evolved  through  the 
development  and  combination  of  the  simpler  ones.  It  has  been 
urged  a_gainst  the  views  here  maintained,  that  it  is  impossible 
to  say  at  what  point  in  the  ascending  scale  animals  become 
capable  of  abstraction,  &c.;  but  who  can  say  at  what  age  this 
occurs  in  our  young  children?  We  see  at  least  that  such  powers 
are  developed  in  children  by  imperceptible  degrees. 

That  animals  retain  their  mental  individuality  is  unquestion- 
able. When  my  voice  awakened  a  train  of  old  associations  in 
the  mind  of  the  before-mentioned  dog,  he  must  have  retained 
his  mental  Individuality,  although  every  atom  of  his  brain  had 

^  Mr.  Hookham,  in  a  letter  to  Prof.  Max  Muller,  in  the  'Birmingham 
News,'  May,  1873. 

*^  'Conferences  sur  la  Theorie  Darwinienne,'  French  translat.  1869, 
p.  132. 


a2  THE  DESCENT  OF  MAN. 

probably  undergone  change  more  than  once  during  the  interval 
of  five  years.  This  dog  might  have  brought  forward  the  argu- 
ment lately  advanced  to  crush  all  evolutionists,  and  said,  "I  abide 

"amid  all  mental  moods  and  all  material  changes The  teaching 

"that  atoms  leave  their  impressions  as  legacies  to  other  atoms 
"falling  into  the  places  they  have  vacated  is  contradictory  of  the 
"utterance  of  consciousness,  and  is  therefore  false;  but  it  is  the 
"teaching  necessitated  by  evolutionism,  consequently  the  hypothe- 
"sis  is  a  false  one."*" 

Language.  — This  faculty  has  justly  been  considered  as  one  of 
the  chief  distinctions  between  man  and  the  lower  animals.  But 
man,  as  a  highly  competent  judge.  Archbishop  Whately  remarks, 
"is  not  the  only  animal  that  can  make  use  of  language  to  express 
"what  is  passing  in  his  mind,  and  can  understand,  more  or  less, 
"what  is  so  expressed  by  another."*^  In  Paraguay  the  Cebus 
azarse  when  excited  utters  at  least  six  distinct  sounds,  which 
excite  in  other  monkeys  similar  emotions.*^  The  movements  of 
the  features  and  gestures  of  monkeys  are  understood  by  us,  and 
they  partly  understand  ours,  as  Rengger  and  others  declare.  It 
is  a  more  remarkable  fact  that  the  dog,  since  being  domesticated, 
has  learnt  to  bark*^  in  at  least  four  or  five  distinct  tones.  Al- 
though barking  is  a  new  art,  no  doubt  the  wild  parent-species  of 
the  dog  expressed  their  feelings  by  cries  of  various  kinds.  With 
the  domesticated  dog  we  have  the  bark  of  eagerness,  as  in  the 
chase;  that  of  anger,  as  well  as  growling;  the  yelp  or  hov/1  of 
despair,  as  when  shut  up;  the  baying  at  night;  the  bark  of  joy,  as 
when  starting  on  a  walk  with  his  master;  and  the  very  distinct 
one  of  demand  or  supplication,  as  when  wishing  for  a  door  or 
window  to  be  opened.  According  to  Houzeau,  who  paid  particular 
attention  to  the  subject,  the  domestic  fowl  utters  at  least  a  dozes 
significant  sounds.^" 

The  habitual  use  of  articulate  language  is,  however,  peculiai 
to  man;  but  he  uses,  in  common  with  the  lower  animals,  inarticu- 
late cries  to  express  his  meaning,  aided  by  gestures  and  the  move. 
ments  of  the  muscles  of  the  face.''  This  especially  holds  good 
with  the  more  sim^ple  and  vivid  feelings,  which  are  but  little 
connected  with  our  higher  intelligence.    Our  cries  of  pain,  fear 


^  The  Rev.  Dr.  J.  McCann,  'Anti-Darwinism,'  1869,  p.  13. 

^7  Quoted  in  'Anthropological  Review,'  1864,  p.  158. 

*^  Reng-g-er,  ibid.  s.  45. 

*9  See  my  'Variation  of  Animals  and  Plants  under  Domestication, 
vol.  i.  p.  27. 

™  'Facultes  Mentales  des  Animaux,'  torn.  ii.  1872,  p.  346-349. 

"1  See  a  discussion  on  this  subject  in  Mr.  E.  B.  Tylor's  very  interest- 
ing work,  'Researches  into  the  Early  History  of  Mankind,'  1S65,  chaps, 
ii.  to  iv. 


MENTAL    POWERS.  83 

surprise,  anger,  together  with  their  appropriate  actions,  and  the 
murmur  of  a  mother  to  her  beloved  child,  are  more  expressive 
than  any  words.  That  which  distinguishes  man  from  the  lower 
animals  is  not  the  understanding  of  articulate  sounds,  for,  as 
every  one  knov»rs,  dogs  understand  many  words  and  sentences. 
In  this  respect  they  are  at  the  same  stage  of  development  as 
infants,  betv/een  the  ages  of  ten  and  twelve  months,  v/ho  under- 
stand many  words  and  short  sentences,  but  cannot  yet  utter  a 
single  word.  It  is  not  the  mere  articulation  which  is  our  dis- 
tinguishing character,  for  parrots  and  other  birds  possess  this 
power.  Nor  is  it  the  mere  capacity  of  connecting  definite  sounds 
with  definite  ideas;  for  it  is  certain  that  some  parrots,  which  have 
been  taught  to  speak,  connect  unerringly  words  with  things,  and 
persons  with  events.^^  The  lov/er  animals  differ  from  man  solely 
in  his  almost  infinitely  larger  power  of  associating  together  the 
most  diversified  sounds  and  ideas;  and  this  obviously  depends  on 
the  high  development  of  his  mental  powers. 

As  Home  Took,  one  of  the  founders  of  the  noble  science  of 
philology,  observes,  language  is  an  art,  like  brewing  or  baking; 
but  writing  would  have  been  a  better  simile.  It  certainly  is  not 
a  true  instinct,  for  every  language  has  to  be  learnt.  It  differs, 
however,  widely  from  all  ordinary  arts,  for  man  has  an  instinctive 
tendency  to  speak,  as  we  see  in  the  babble  of  our  young  children; 
whilst  no  child  has  an  instinctive  tendency  to  brew,  bake,  or 
write.  Moreover,  no  philologist  now  supposes  that  any  language 
has  been  deliberately  invented;  it  has  been  slowly  and  uncon- 
sciously developed  by  many  steps.^^    The  sounds  uttered  by  birds 

52  I  have  received  several  detailed  accounts  to  this  effect.  Admiral 
Sir  J.  Sullivan,  whom  I  know  to  be  a  careful  observer,  assures  me  that 
an  African  parrot,  long  kept  in  his  father's  house,  invariably  called 
certain  persons  of  the  household,  as  well  as  visitors,  by  their  names. 
He  said  "good  morning"  to  every  one  at  breakfast,  and  "good  night" 
to  each  as  they  left  the  room  at  night,  and  never  reversed  these  salu- 
tations. To  Sir  J.  Sullivan's  father  lie  used  to  add  to  the  "good  morn- 
ing" a  short  sentence,  which  was  never  once  repeated  after  his  father's 
death.  He  scolded  violently  a  strange  dog  which  came  into  the  room 
through  the  open  window;  and  he  scolded  another  parrot  (saying 
"you  naughty  polly")  which  had  got  out  of  its  cage,  and  v/as  eating 
apples  on  the  kitchen  table.  See  also,  to  the  same  effect,  Houzeau  en 
parrots,  'Facultes  Mentales,'  torn.  ii.  p.  309.  Dr.  A.  Moschkan  informs 
me  that  he  knew  a  starling  which  never  made  a  mistake  in  saying  in 
German  "good  morning"  to  persons  arriving,  and  "good-bye,  old  fel- 
low," to  those  departing.    I  could  add  several  other  such  cases. 

53  See  some  good  remarks  on  this  head  by  Prof.  Whitney,  in  his 
•Oriental  and  Linguistic  Studies,'  1873,  p.  354.  He  observes  that  the 
desire  of  communication  between  man  is  the  living  force,  which,  in 
the  development  of  language,    "works  both  consciously  ajid  uncon- 


84  THE  DESCENT  OF  MAN. 

offer  in  several  respects  th.e  nearest  analogy  to  language,  for  all 
the  members  of  the  same  species  utter  the  same  instinctive  cries 
expressive  of  their  emotions;  and  all  the  kinds  which  sing,  exert 
their  power  instinctively;  but  the  actual  song,  and  even  the  call 
notes,  are  learnt  from  their  parents  or  foster-parents.  These 
sounds,  as  Daines  Barrington^*  has  proved,  "are  no  more  innate 
"than  language  is  in  man,"  The  first  attempts  to  sing  "may  be 
"compared  to  the  imperfect  endeavor  in  a  child  to  babble."  The 
young  males  continue  practicing,  or  as  the  bird-catchers  say, 
"recording,"  for  ten  or  eleven  months.  Their  first  essays  show 
hardly  a  rudiment  of  the  future  song;  but  as  they  grow  older 
we  can  perceive  what  they  are  aiming  at;  and  at  last  they  are 
said  "to  sing  their  song  round."  Nestlings  which  have  learnt  the 
song  of  a  distinct  species,  as  with  the  canary-birds  educated  in  the 
Tyrol,  teach  and  transmit  their  new  song  to  their  offspring.  The 
slight  natural  differences  of  song  in  the  same  species  inhabiting 
different  districts  may  be  appositely  compared,  as  Barrington 
remarks,  "to  provincial  dialects;"  and  the  songs  are  allied,  though 
distinct  species  may  be  compared  with  the  languages  of  distinct 
races  of  man.  I  have  given  the  foregoing  details  to  show  that 
an  instinctive  tendency  to  acquire  an  art  is  not  peculiar  to  man. 

With  respect  to  the  origin  of  articulate  language,  after  having 
read  on  the  one  side  the  highly  interesting  works  of  Mr.  Hens- 
leigh  Wedgwood,  the  Rev.  F.  Parrar,  and  Prof.  Schleicher,^"  and 
the  celebrated  lectures  of  Prof.  Max  Miiller  on  the  other  side,  I 
cannot  doubt  that  language  owes  its  origin  to  the  imitation  and 
modification  of  various  natural  sounds,  the  voices  of  other  ani- 
mals, and  man's  own  instinctive  cries,  aided  by  signs  and  gestures. 
When  we  treat  of  sexual  selection  we  shall  see  that  primeval  man, 
or  rather  some  early  progenitor  of  man,  probably  first  used  his 
voice  in  producing  true  musical  cadences,  that  is  in  singing,  as 
do  some  of  the  gibbon-apes  at  the  present  day;  and  we  may  con- 
clude from  a  widely-spread  analogy,  that  this  power  would  have 
been  especially  exerted  during  the  courtship  of  the  sexes, — would 
have  expressed  various  emotions,  such  as  love,  jealousy,  triumph, 
— and  would  have  served  as  a  challenge  to  rivals.    It  is,  therefore, 

"sciously;    consciously  as  reg-ards  the  immediate  end  to  be  attained; 
"unconsciously  as  regards  the  further  consequences  of  the  act." 

54  Hon.  Daines  Barring-ton  in  'Philosoph.  Transactions,'  1773,  p.  262. 
See  also  Dureau  de  la  Malle,  in  'Ann.  des.  Sc.  Nat.'  3rd  series,  Zoolog-. 
tom.  X.  p.  119. 

55  'On  the  Origin  of  Language,'  by  H.  Wedgwood,  1866.  'Chapters  on 
Language,'  by  the  Rev.  P.  W.  Farrar,  1865.  These  works  are  most  in- 
teresting. See  also  'De  la  Phys.  et  de  Parole'  par  Albert  Lemoine,  1865, 
p.  190.  The  work  on  this  subject,  by  the  late  Prof.  Aug.  Schleicher, 
has  been  translated  by  Dr.  Bikkers  into  English,  under  the  title  of 
'Darwinism  tested  by  the  Science  of  Language,'  1869. 


MENTAL    POWERS.  85 

probable  that  the  imitation  of  musical  cries  by  articulate  sounds 
may  have  given  rise  to  words  expressive  of  various  complex 
emotions.  The  strong  tendency  in  our  nearest  allies,  the  monkeys, 
in  microcephalous  idiots,^"  and  in  the  barbarous  races  of  mankind, 
to  imitate  v?-hatever  they  hear  deserves  notice,  as  bearing  on  the 
subject  of  imitation.  Since  monkeys  certainly  understand  much 
that  is  said  to  them  by  man,  and  when  wild,  utter  signal-cries  of 
danger  to  their  fellows;"  and  since  fowls  give  distinct  warnings 
for  danger  on  the  ground,  or  in  the  sky  from  hawks  (both,  as 
well  as  a  third  cry,  intelligible  to  dogs)  ,^^  may  not  some  unusually 
wise  ape-like  animal  have  imitated  the  growl  of  a  beast  of  prey, 
and  thus  told  his  fellow-monkeys  the  nature  of  the  expected 
danger?  This  would  have  been  a  first  step  in  the  formation  of  a 
language. 

As  the  voice  was  used  more  and  more,  the  vocal  organs  would 
have  been  strengthened  and  perfected  through  the  principle  of 
the  inherited  effects  of  use;  and  this  would  have  reacted  on  the 
power  of  speech.  But  the  relation  between  the  continued  use  of 
language  and  the  development  of  the  brain,  has  no  doubt  been 
far  more  important.  The  mental  powers  in  some  early  progenitor 
of  man  must  have  been  more  highly  developed  than  in  any  existing 
ape,  before  even  the  most  imperfect  form  of  speech  could  have 
come  into  use;  but  we  may  confidently  believe  that  the  con- 
tinued use  and  advancement  of  this  power  would  have  reacted  on 
the  mind  itself,  by  enabling  and  encouraging  it  to  carry  on  long 
trains  of  thought.  A  complex  train  of  thought  can  no  more  be 
carried  on  without  the  aid  of  words,  whether  spoken  or  silent, 
than  a  long  calculation  without  the  use  of  figures  or  algebra.  It 
appears,  also,  that  even  an  ordinary  train  of  thought  almost  re- 
quires, or  is  greatly  facilitated  by  some  form  of  language,  for 
the  dumb,  deaf,  and  blind  girl,  Laura  Bridgman,  was  observed  to 
use  her  fingers  whilst  dreaming.^^  Nevertheless,  a  long  succession 
of  vivid  and  connected  ideas  may  pass  through  the  mind  without 
the  aid  of  any  form  of  language,  as  we  may  infer  from  the  move- 
ments of  dogs  during  their  dreams.  We  have,  also,  seen  that  ani- 
mals are  able  to  reason  to  a  certain  extent,  manifestly  without 
the  aid  of  language.  The  intimate  connection  between  the  brain, 
as  it  is  now  developed  in  us,  and  the  faculty  of  speech,  is  well 


s«  Vog-t,  'Memoire  sur  les  Microcephales,'  1867,  p.  169.  With  respect  to 
savag-es,  I  have  given  some  facts  in  my  'Journal  of  Researches,'  &c., 
1845,  p.  206. 

^'^  See  clear  evidence  on  this  head  in  the  two  v/crks  so  often  quoted, 
by  Brehm  and  Rengger. 

^  Houzeau  gives  a  very  curious  account  of  his  observations  on  this 
subject  in  his  'Facultes  Mentales  des  Animaux,'  tom.  ii.,  p.  348. 

50  See  remarks  on  this  head  by  Dr.  Maudsley,   'The  Physiology  and 
Pathology  of  Mind,'  2nd  edit.  1868,  p.  199. 
7 


gg  THE  DESCENT  OF  MAN. 

shown  by  those  curious  cases  of  brain-disease  in  which  speech  is 
specially  affected,  as  when  the  power  to  remember  substantives 
is  lost,  whilst  other  words  can  be  correctly  used,  or  where  sub- 
stantives of  a  certain  class,  or  all  except  the  initial  letters  of 
substantives  and  proper  names  are  forgotten.*^  There  is  no  more 
improbability  in  the  continued  use  of  the  mental  and  vocal  organs 
leading  to  inherited  changes  in  their  structure  and  functions 
than  in  the  case  of  handwriting,  which  depends  partly  on  the 
form  of  the  hand  and  partly  on  the  disposition  of  the  mind;  and 
hand-writing  is  certainly  inherited.®^ 

Several  writers,  more  especially  Prof.  Max  Miiller,*'-  have  lately 
insisted  that  the  use  of  language  implies  the  power  of  forming 
general  concepts;  and  that  as  no  animals  are  supposed  to  possess 
this  power,  an  impossible  barrier  is  formed  between  them  and 
man.^  With  respect  to  animals,  I  have  already  endeavored  to 
show  that  they  have  this  power,  at  least  in  a  rude  and  incipient 
degree.  As  far  as  concerns  infants  of  from  ten  to  eleven  months 
old,  and  deaf-mutes,  it  seems  to  me  incredible,  that  they  should 
be  able  to  connect  certain  sounds  with  certain  general  ideas  as 
quickly  as  they  do,  unless  such  ideas  were  already  formed  in 
their  minds.  The  same  remark  may  be  extended  to  the  more  in- 
telligent animals;  as  Mr.  Leslie  Stephen  observes/*  "A  dog 
"frames  a  general  concept  of  cats  or  sheep,  and  knows  the  cor- 

^  Many  curious  cases  have  been  recorded.  See,  for  instance,  Dr. 
Bateman  'On  Aphasia,'  1870,  pp.  27,  31,  53,  100,  &c.  Also,  'Inquiries  Con- 
cerning the  Intellectual  Powers,'  by  Dr.  Abercrombie,  1838,  p.  150. 

«i  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
ii.  p.  6. 

«2  Lectures  on  'Mr.  Darwin's  Philosophy  of  Lang-uag-e,'  1873. 

«8  The  judgment  of  a  distinguished  philologist,  such  as  Prof.  Whitney, 
will  have  far  more  weight  on  this  point  than  anything  that  I  can  say. 
He  remarks  ('Oriental  and  Linguistic  Studies,'  1873,  p.  297),  in  speaking 
of  Bleek's  views:  "Because  on  the  grand  scale  language  is  the  neces- 
"sary  auxiliary  of  thought,  indispensable  to  the  development  of  the 
"power  of  thinking,  to  the  distinctness  and  variety  and  complexity  of 
"cognitions  to  the  full  mastery  of  consciousness;  therefore  he  would 
"fain  make  thought  absolutely  impossible  without  speech,  identifying 
"the  faculty  with  its  instrument.  He  might  just  as  reasonably  assert 
"that  the  human  hand  cannot  act  without  a  tool.  With  such  a  doctrine 
"to  start  from,  he  cannot  stop  short  of  Muller's  worst  paradoxes,  that 
"an  infant  (in  fans,  not  speaking)  is  not  a  human  being,  and  that  deaf 
"mutes  do  not  become  possessed  of  reason  until  they  learn  to  twist 
"their  fingers  into  imitation  of  spoken  words."  Max  Muller  gives  in 
italics  ('Lectures  on  Mr.  Darwin's  Philosophy  of  Language,'  1873,  th'rd 
lecture)  the  following  aphorism:  "There  is  no  thought  without  words, 
"as  little  as  there  are  words  without  thought."  What  a  strange  defi- 
nition must  here  be  given  to  the  word  thought! 
«  'Essays  on  Free- thinking,'  &c.,  1873,  p.  82. 


MENTAL   POWERS.  87 

"responding  words  as  well  as  a  philcsoptier.  And  the  capacity  to 
"understand  is  as  good  a  proof  of  vocal  intelligence,  though  in  an 
"inferior  degree,  as  the  capacity  to  speak." 

Why  the  organs  now  used  for  speech  should  have  heen  originally 
perfected  for  this  purpose,  rather  than  any  other  organs,  it  is 
not  difficult  to  see.  Ants  have  considerable  powers  of  intercom- 
munication by  means  of  their  antennae,  as  shown  by  Huber,  who 
devotes  a  whole  chapter  to  their  language.  We  might  have  used 
our  fingers  as  efficient  instruments,  for  a  person  with  practice 
can  report  to  a  deaf  man  every  word  of  a  speech  rapidlj'-  delivered 
at  a  public  meeting;  but  the  loss  of  our  hands,  whilst  thus  em- 
ployed, would  have  been  a  serious  inconvenience.  As  all  the 
higher  mammals  possess  vocal  organs,  constructed  on  the  same 
general  plan  as  ours,  and  used  as  a  means  of  communication,  it 
was  obviously  probable  that  these  same  organs  would  be  still  fur- 
ther developed  if  the  power  of  communication  had  to  be  im- 
proved; and  this  has  been  effected  by  the  aid  of  adjoining  and 
well  adapted  parts,  namely  the  tongue  and  lips.*^^  The  fact  ^of 
the  higher  apes  not  using  their  vocal  organs  for  speech,  no  doubt 
depends  on  their  intelligence  not  having  been  sufficiently  ad- 
vanced. The  possession  by  them  of  organs,  which  with  long- 
continued  practice  might  have  been  used  for  speech,  although 
not  thus  used,  is  paralleled  by  the  case  of  many  birds  which  pos- 
sess organs  fitted  for  singing,  though  they  never  sing.  Thus,  the 
nightingale  and  crow  have  vocal  organs  similarly  constructed, 
these  being  used  by  the  former  for  diversified  song,  and  by  the 
latter  only  for  croaking.*^®  If  it  be  asked  why  apes  have  not  had 
their  intellects  developed  to  the  same  degree  as  that  of  man, 
general  causes  only  can  be  assigned  in  answer,  and  it  is  unreason- 
able to  expect  anything  more  definite,  considering  our  ignorance 
with  respect  to  the  successive  stages  of  development  through 
which  each  creature  has  passed. 

The  formation  of  different  languages  and  of  distinct  species, 
and  the  proofs  that  both  have  been  developed  through  a  gradual 
process,  are  curiously  parallel."    But  we  can  trace  the  formation 

«5  See  some  good  remarks  to  this  effect  by  Dr.  Maudsley,  'The  Phys- 
iology and  Pathology  of  Mind,'    1868,  p.  199. 

66  Macgillivray,  'Hist,  of  British  Birds,'  vol.  ii.  1839,  p.  29.  An  excel- 
lent observer,  Mr.  Blackwall  remarks  that  the  magpie  learns  to  pro- 
nounce single  words,  and  even  short  sentences,  more  readily  than 
almost  any  other  British  bird;  yet,  as  he  adds,  after  long  and  closely 
investigating  its  habits,  he  has  never  known  it  in  a  state  of  nature, 
display  any  unusual  capacity  for  imitation.  'Researches  in  Zoology,' 
1834,  p.  158. 

6'^  See  the  very  interesting  parallelism  between  the  development  of 
species  and  languages,  given  by  Sir  C.  Lyell  in  'The  Geolog.  Evidences 
of  the  Antiquity  of  Man,'  1863,  chap,  xxiii. 


V^ 


88  THE  DESCENT  OF  MAN. 

of  many  words  further  back  than  that  of  species,  for  we  can 
perceive  how  they  actually  arose  from  the  imitation  of  various 
sounds.  We  find  in  distinct  languages  striking  homologies  due 
to  community  of  descent,  and  analogies  due  to  a  similar  process 
of  formation.  The  manner  in  which  certain  letters  or  sounds 
change  when  others  change  is  very  like  correlated  growth.  We 
have  in  both  cases  the  reduplication  of  parts,  the  effects  of  long- 
continued  use,  and  so  forth.  The  frequent  presence  of  rudi- 
ments, both  in  language  and  in  species,  is  still  more  remarkable. 
The  letter  m  in  the  word  am,  means  I;  so  that  in  the  expression 
I  am,  a  superfluous  and  useless  rudiment  has  been  retained.  In 
the  spelling  also  of  words,  letters  often  remain  as  the  rudiments 
of  ancient  forms  of  pronunciation.  Languages,  like  organic  be- 
ings, can  be  classed  in  groups  under  groups;  and  they  can  be 
classed  either  naturally  according  to  descent,  or  artificially  by 
other  characters.  Dominant  languages  and  dialects  spread  widely, 
and  lead  to  the  gradual  extinction  of  other  tongues.  A  language, 
like  a  species,  when  once  extinct,  never,  as  Sir  C.  Lyell  remarks, 
reappears.  The  same  language  never  has  two  birth-places.  Dis- 
tinct languages  may  be  crossed  or  blended  together.^^  We  see 
variability  in  every  tongue,  and  new  words  are  continually  crop- 
ping up;  but  as  there  is  a  limit  to  the  powers  of  the  memory, 
single  words,  like  whole  languages,  gradually  become  extinct.  As 
Max  Miiller^^  has  well  remarked: — "A  struggle  for  life  is  con- 
"stantly  going  on  amongst  the  words  and  grammatical  forms  in 
"each  language.  The  better,  the  shorter,  the  easier  forms  are  con- 
"stantly  gaining  the  upper  hand,  and  they  owe  their  success  to 
"their  own  inherent  virtue."  To  these  more  important  causes 
of  the  survival  of  certain  words,  mere  novelty  and  fashion  may  be 
added;  for  there  is  in  the  mind  of  man  a  strong  love  for  slight 
changes  in  all  things.  The  survival  or  preservation  of  certain 
favored  words  in  the  struggle  for  existence  is  natural  selection. 

The  perfectly  regular  and  wonderfully  complex  construction 
of  the  languages  of  many  barbarous  nations  has  often  been  ad- 
vanced as  a  proof,  either  of  the  divine  origin  of  these  languages, 
or  of  the  high  art  and  former  civilization  of  their  founders.  Thus 
F.  von  Schlegel  writes:  "In  those  languages  which  appear  to 
"be  at  the  lowest  grade  of  intellectual  culture,  we  frequently 
"observe  a  very  high  and  elaborate  degree  of  art  in  their  gram- 
"matical  structure.  This  is  especially  the  case  with  the  Basque 
"and  the  Lapponian,  and  many  of  the  American  languages."^"  But 

«8  See  remarks  to  this  effect  by  the  Rev.  F.  W.  Farrar,  in  an  inter- 
esting- article,  entitled  'Philology  and  Darwinism'  in  'Nature,'  March 
24th,  1870,  p.  528. 

«»  'Nature,'  Jan.  6th,  1870,  p.  257. 

™  Quoted  by  C.  S.  Wake,  'Chapters  on  Man,'  1868,  p.  lOL 


MENTAL    POWERS.  '  89 

it  is  assuredly  an  error  to  speak  of  any  language  as  an  art,  in  the 
sense  of  its  having  been  elaborately  and  methodically  formed. 
Philologists  now  admit  that  conjugations,  declensions,  &c.,  orig- 
inally existed  as  distinct  words,  since  joined  together;  and  as 
such  words  express  the  most  obvious  relations  between  objects 
and  persons,  it  is  not  surprising  that  they  should  have  been  used 
by  the  men  of  most  races  during  the  earliest  ages.  With  respect 
to  perfection,  the  following  illustration  will  best  show  how  easily 
we  may  err;  a  Crinoid  sometimes  consists  of  no  less  than  150,000 
pieces  of  shell,^^  all  arranged  with  perfect  symmetry  in  radiating 
lines;  but  a  naturalist  does  not  consider  an  animal  of  this  kind 
as  more  perfect  than  a  bilateral  one  with  comparatively  few 
parts,  and  with  none  of  these  parts  alike,  excepting  on  the  op- 
posite sides  of  the  body.  He  justly  considers  the  differentiation 
and  specialization  of  organs  as  the  test  of  perfection.  So  with 
languages;  the  most  symmetrical  and  complex  ought  not  to  be 
ranked  above  irregular,  abbreviated,  and  bastardized  languages, 
which  have  borrowed  expressive  words  and  useful  forms  of  con- 
struction from  various  conquering,  conquered,  or  immigrant  races. 
From  these  few  and  imperfect  remarks  I  conclude  that  the  ex- 
tremely complex  and  regular  construction  of  many  barbarous  lan- 
guages, is  no  proof  that  they  owe  their  origin  to  a  special  act 
of  creation.'-  Nor,  as  we  have  seen,  does  the  faculty  of  articulate 
speech  in  itself  offer  any  insuperable  objection  to  the  belief  that 
man  has  been  developed  from  some  lower  form. 

Sense  of  Beauty.  — This  sense  has  been  declared  to  be  peculiar 
to  man.  I  refer  here  only  to  the  pleasure  given  by  certain  colors, 
forms,  and  sounds,  and  which  may  fairly  be  called  a  sense  of  the 
beautiful;  with  cultivated  men  such  sensations  are,  hov/ever, 
intimately  associated  with  complex  ideas  and  trains  of  thought. 
V/hen  we  behold  a  male  bird  elaborately  displaying  his  graceful 
plumes  or  splendid  colors  before  the  female,  whilst  other  birds, 
not  thus  decorated,  make  no  such  display,  it  is  impossible  to 
doubt  that  she  admires  the  beauty  of  her  male  partner.  As 
women  everywhere  deck  themselves  with  these  plumes,  the  beauty 
of  such  ornaments  cannot  be  disputed.  As  we  shall  see  later, 
the  nests  of  humming-birds,  and  the  playing  passages  of  bower- 
birds  are  tastefully  ornamented  with  gayly-colored  objects;  and 
this  shows  that  they  must  receive  some  kind  of  pleasure  from  the 
sight  of  such  things.  Yv^ith  the  great  majority  of  animals,  how- 
ever, the  taste  for  the  beautiful  is  confined,  as  far  as  we  can  judge, 
to  the  attractions  of  the  opposite  sex.    The  sweet  strains  poured 

■^1  Buckland,  'Bridgewater  Treatise,'  p.  411. 

■^2  See  some  g-ood  remarks  on  the  simplification  of  languages,  hy  Sir  J. 
Lubbock,  'Origin  of  Civilization,'  1870,  p.  278. 


^ 


90  THE  DESCENT  OF  MAN. 

forth  by  many  male  birds  during  the  season  of  love,  are  certainly 
admired  by  the  females,  of  which  fact  evidence  will  hereafter  be 
given.  If  female  birds  had  been  incapable  of  appreciating  the 
beautiful  colors,  the  ornaments,  and  voices  of  their  male  partners, 
all  the  labor  and  anxiety  exhibited  by  the  latter  in  displaying  their 
charms  before  the  females  would  have  been  thrown  away;  and 
this  it  is  impossible  to  admit.  Why  certain  bright  colors  should 
excite  pleasure  cannot,  I  presume,  be  explained,  any  more  than 
why  certain  flavors  and  scents  are  agreeable;  but  habit  has 
something  to  do  with  the  result,  for  that  which  is  at  first  un- 
pleasant to  our  senses,  ultimately  becomes  pleasant,  and  habits 
are  inherited.  With  respect  to  sounds,  Helmholtz  has  explained 
to  a  certain  extent  on  physiological  principles,  why  harmonies 
and  certain  cadences  are  agreeable.  But  besides  this,  sounds  fre- 
quently recurring  at  irregular  intervals,  are  highly  disagreeable, 
as  every  one  will  admit  who  has  listened  at  night  to  the  irregular 
flapping  of  a  rope  on  board  ship.  The  same  principle  seems  to 
come  into  play  with  vision,  as  the  eye  prefers  symmetry  or  figures 
with  some  regular  recurrence.  Patterns  of  this  kind  are  em- 
ployed by  even  the  lowest  savages  as  ornaments;  and  they  have 
been  developed  through  sexual  selection  for  the  adornment  of 
some  male  animals.  Whether  we  can  or  not  give  any  reason  for 
the  pleasure  thus  derived  from  vision  and  hearing,  yet  man  and 
many  of  the  lower  animals  are  alike  pleased  by  the  same  colors, 
graceful  shading  and  forms,  and  the  same  sounds. 

The  taste  for  the  beautiful,  at  least  as  far  as  female  beauty  is 
concerned,  is  not  of  a  special  nature  in  the  human  mind;  for  it 
differs  widely  in  the  different  races  of  man,  and  is  not  quite  the 
same  even  in  the  different  nations  of  the  same  race.  Judging 
from  the  hideous  ornaments,  and  the  equally  hideous  music  ad- 
mired by  most  savages,  it  might  be  urged  that  their  aesthetic 
faculty  was  not  so  highly  developed  as  in  certain  animals,  for 
instance,  as  in  birds.  Obviously  no  animal  would  be  capable  of 
admiring  such  scenes  as  the  heavens  at  night,  a  beautiful  land- 
scape, or  refined  music;  but  such  high  tastes  are  acquired  through 
culture,  and  depend  on  complex  associations;  they  are  not  en- 
joyed by  barbarians  or  by  uneducated  persons. 

Many  of  the  faculties,  which  have  been  of  inestimable  service 
to  man  for  his  progressive  advancement,  such  as  the  powers  of 
the  imagination,  wonder,  curiosity,  an  undefined  sense  of  beauty, 
a  tendency  to  imitation,  and  the  love  of  excitement  or  novelty, 
could  hardly  fail  to  lead  to  capricious  changes  of  customs  and 
fashions.  I  have  alluded  to  this  point,  because  a  recent  writer'^ 
has  oddly  fixed  on  Caprice  "as  one  of  the  most  remarkable  and 
"typical  differences  between  savages  and  brutes."     But  not  only 

'3  'The  Spectator,'  Dec.  4th  1869,  p.  1430. 


MENTAL.    POWERS.  91 

can  we  partially  understand  how  it  is  that  man  is  from  various 
conflicting  influences  rendered  capricious,  but  that  the  lower 
animals  are,  as  we  shall  hereafter  see,  likewise  capricious  in  their 
affections,  aversions,  and  sense  of  beauty.  There  is  also  reason 
to  suspect  that  they  love  novelty,  for  its  own  sake. 

Belief  in  Ood — Religion. — There  is  no  evidence  that  man  was 
aboriginally  endowed  with  the  ennobling  belief  in  the  existence 
of  an  Omnipotent  God.  On  the  contrary  there  is  ample  evidence, 
derived  not  from  hasty  travelers,  but  from  men  who  have  long  »/ 
resided  with  savages,  that  numerous  races  have  existed,  and  still 
exist,  who  have  no  idea  of  one  or  more  gods,  and  who  have  no 
words  in  their  languages  to  express  such  an  idea.'^*  The  question 
is  of  course  wholly  distinct  from  that  higher  one,  whether  there 
exists  a  Creator  and  Ruler  of  the  universe;  and  this  has  been 
answered  in  the  affirmative  by  some  of  the  highest  intellects  that 
have  ever  existed. 

If,  however,  we  include  under  the  term  "religion"  the  belief 
in  unseen  or  spiritual  agencies,  the  case  is  wholly  different;  for 
this  belief  seems  to  be  universal  with  the  less  civilized  races. 
Nor  is  it  difficult  to  comprehend  how  it  arose.  As  soon  as  the 
important  faculties  of  the  imagination,  wonder,  and  curiosity, 
together  with  some  power  of  reasoning,  had  become  partially 
developed,  man  would  naturally  crave  to  understand  what  was 
passing  around  him,  and  would  have  vaguely  speculated  on  his 
own  existence.  As  Mr.  M'Lennan'^^  has  remarked,  "Some  explana- 
"tion  of  the  phenomena  of  life,  a  man  must  feign  for  himself; 
"and  to  judge  from  the  universality  of  it,  the  simplest  hypothesis, 
"and  the  first  to  occur  to  men,  seems  to  have  been  that  natural 
"phenomena  are  ascribable  to  the  presence  in  animals,  plants, 
"and  things,  and  in  the  forces  of  nature,  of  such  spirits  prompting 
"to  action  as  men  are  conscious  they  themselves  possess."  It 
is  also  probable,  as  Mr.  Tylor  has  shown,  that  dreams  may  have 
first  given  rise  to  the  notion  of  spirits;  for  savages  do  not  readily 
distinguish  between  subjective  and  objective  impressions.  When 
a  savage  dreams,  the  figures  which  appear  before  him  are  be- 
lieved to  have  come  from  a  distance,  and  to  stand  over  him;  or 
"the  soul  of  the  dreamer  goes  out  on  its  travels,  and  comes  home 
"with  a  remembrance  of  what  it  has  seen."^^    But  until  the  facul- 

'*  See  an  excellent  article  on  this  subject  by  the  Rev.  F.  W.  Farrar, 
in  the  'Anthropological  Review,'  Aug-.,  1864,  p.  ccxvii.  For  further 
facts  see  Sir  J.  Lubbock,  'Prehistoric  Times,'  2nd  edit.  1869,  p.  564;  and 
especially  the  chapters  on  Religion  in  his  'Origin  of  Civilization,'  1870. 

"^^  'The  Worship  of  Animals  and  Plants,'  in  the  'Fortnightly  Review,' 
Oct.  1,  1869,  p.  422. 

"s  Tyior,  'Early  History  of  Mankind,'  1865,  p.  6.  See  also  the  three 
striking  chapters  on  the  Development  of  Religion,  in  Lubbock's  'Origin 


92  THE   DESCENT  OF  MAN. 

ties  of  imagination,  curiosity,  reason,  «fec.,  had  been  fairly  well 
developed  in  the  mind  of  man,  his  dreams  would  not  have  led  him 
to  believe  in  spirits,  any  more  than  in  the  case  of  a  dog. 

The  tendency  in  savages  to  imagine  that  natural  objects  and 
agencies  are  animated  by  spiritual  or  living  essences,  is  perhaps 
illustrated  by  a  little  fact  which  I  once  noticed:  my  dog,  a  full- 
grown  and  very  sensible  animal,  was  lying  on  the  lawn  during  a 
hot  and  still  day; '  but  at  a  little  distance  a  slight  breeze  occasion- 
ally moved  an  open  parasol,  which  would  have  been  wholly  dis- 
regarded by  the  dog,  had  any  one  stood  near  it.  As  it  was,  every 
time  that  the  parasol  slightly  moved,  the  dog  growled  fiercely 
and  barked.  He  must,  I  think,  have  reasoned  to  himself  in  a 
rapid  and  unconscious  manner,  that  movement  without  any  ap- 
parent cause  indicated  the  presence  of  some  strange  living  agent, 
and  that  no  stranger  had  a  right  to  be  on  his  territory. 

The  belief  in  spiritual  agencies  would  easily  pass  into  the 
belief  in  the  existence  of  one  or  more  gods.  For  savages  would 
naturally  attribute  to  spirits  the  same  passions,  the  same  love  of 
vengeance  or  simplest  form  of  justice,  and  the  same  affections 
which  they  themselves  feel.  The  Fuegians  appear  to  be  in  this 
respect  in  an  intermediate  condition,  for  when  the  surgeon  on 
board  the  "Beagle"  shot  some  young  ducklings  as  specimens,  York 
Minster  declared  in  the  most  solemn  manner,  "Oh,  Mr.  Bynoe, 
"much  rain,  much  snow,  blow  much;"  and  this  was  evidently  a 
retributive  punishment  for  wasting  human  food.  So  again  he 
related  how,  when  his  brother  killed  a  "wild  man,"  storms  long 
raged,  much  rain  and  snow  fell.  Yet  we  could  never  discover 
that  the  Fuegians  believed  in  what  we  should  call  a  God,  or  prac- 
ticed any  religious  rites;  and  Jemmy  Button,  with  justifiable 
pride,  stoutly  maintained  that  there  was  no  devil  in  his  land. 
This  latter  assertion  is  the  more  remarkable,  as  with  savages  the 
belief  in  bad  spirits  is  far  more  common  than  that  in  good  ones. 

of  Civilization,'  1870.  In  a  like  manner  Mr.  Herbert  Spencer,  in  his 
ingenious  essay  in  the  'Fortnightly  Review'  (May  1st,  1870,  p.  535),  ac- 
counts for  the  earliest  forms  of  religious  belief  throughout  the  world, 
by  man  being  led  through  dreams,  shadows,  and  other  causes,  to  look 
at  himself  as  a  double  essence,  corporeal  and  spiritual.  As  the  spiritual 
being  is  supposed  to  exist  after  death  and  to  be  powerful,  it  is  propi- 
tiated by  various  gifts  and  ceremonies,  and  its  aid  invoked.  He  then 
further  shows  that  names  or  nicknames  given  from  some  animal  or 
other  object,  to  the  early  progenitors  or  founders  of  a  tribe,  are  sup- 
posed after  a  long  interval  to  represent  the  real  progenitor  of  the  tribe; 
and  such  animal  or  object  is  then  naturally  believed  still  to  exist  as 
a  spirit,  is  held  sacred,  and  worshipped  as  a  god.  Nevertheless  I  can- 
not but  suspect  that  there  is  a  still  earlier  and  ruder  stage,  when  any- 
thing which  manifests  power  or  movement  is  thought  to  be  endowed 
with  some  form  of  life,  and  with  mental  faculties  analogous  to  our  own. 


MENTAL   POWERS.  93 

The  feeling  of  religious  devotion  is  a  highly  complex  one,  con- 
sisting of  love,  complete  submission  to  an  exalted  and  mysterious 
superior,  a  strong  sense  of  dependence,^'  fear,  reverence,  gratitude, 
hope  for  the  future,  and  perhaps  other  elements.  No  being  could 
experience  so  complex  an  emotion  until  advanced  in  his  intellec- 
tual and  moral  faculties  to  at  least  a  moderately  high  level.  Never- 
theless, we  see  some  distant  approach  to  this  state  of  mind  in  the 
deep  love  of  a  dog  for  his  master,  associated  with  complete  sub- 
mission, some  fear,  and  perhaps  other  feelings.  The  behavior 
of  a  dog  when  returning  to  his  master  after  an  absence,  and.  as 
I  may  add,  of  a  monkey  to  his  beloved  keeper,  is  widely  different 
from  that  towards  their  fellows.  In  the  latter  case  the  transports 
of  joy  appear  to  be  somewhat  less,  and  the  sense  of  equality  is 
shown  in  every  action.  Professor  Braubach  goes  so  far  as  to 
maintain  that  a  dog  looks  on  his  master  as  on  a  god.^^ 

The  same  high  mental  faculties  which  first  Jed  man  to  believe 
in  unseen  spiritual  agencies,  then  in  fetishism,  polytheism,  and  > 
ultimately  in  monotheism,  would  infallibly  lead  him,  as  long  as 
his^  jeasoning  powers  remained  poorly  developed,  to  various 
"strange  superstitions  and  customs.  Many  of  these  are  terrible 
to  think  of — such  as  the  sacrifice  of  human  beings  to  a  blood-lov- 
ing god;  the  trial  of  innocent  persons  by  the  ordeal  of  poison  or 
fire;  witchcraft,  &c. — yet  it  is  well  occasionally  to  reflect  on  these 
superstitions,  for  they  show  us  what  an  infinite  debt  of  gratitude 
we  owe  to  the  improvement  of  our  reason,  to  science,  and  to  our 
accumulated  knowledge.  As  Sir  J.  Lubbock^''  has  well  observed, 
"it  is  not  too  much  to  say  that  the  horrible  dread  of  unknown  evil 
"hangs  like  a  thick  cloud  over  savage  life,  and  embitters  every 
"pleasure."  These  miserable  and  indirect  consequences  of  our 
highest  faculties  may  be  compared  with  the  incidental  and  occa- 
sional mistakes  of  the  instincts  of  the  lower  animals. 

"  See  an  able  article  on  the  'Physical  Elements  of  Religion,*  by  Mr. 
L.  Owen  Pike,  in  'Anthropolog'.  Review,'  April,  1870,  p.  Ixiii. 

■^8  'Religion,  Moral,  &c.,  der  Darwin'schen  Art-Lehre,'  1869,  s.  53.  It 
is  said  (Dr.  W.  Lauder  Lindsay,  'Journal  of  Mental  Science,'  1871,  p. 
43),  that  Bacon  long  ago,  and  the  poet  Burns,  held  the  same  notion. 

''^  'Prehistoric  Times,'  2nd  edit.  p.  571.  In  this  work  (p.  571)  there  will 
be  found  an  excellent  account  of  the  many  strange  and  capricious  cus- 
toms of  savages. 


94  THE  DESCENT  OF  MAN. 


CHAPTER    IV. 

COMPARISON  OF  THE  MENTAL  POWERS  OF  MAN  AND 
THE  LOWER  ANIMALS— Continued. 

The  moral  sense — Fundamental  proposition — The  qualities  of  social 
animals — Origin  of  sociability — Struggle  between  opposed  instincts — 
Man  a  social  animal — The  more  enduring  social  instincts  conquer 
other  less  persistent  instincts — The  social  virtues  alone  regarded  by 
savages — The  self-regarding  virtues  acquired  at  a  later  stage  of  de- 
velopment—The importance  of  the  judgment  of  the  members  of  the 
same  community  on  conduct— Transmission  of  moral  tendencies- 
Summary. 

I  fully  subscribe  to  the  judgment  of  those  writers^  who  main- 
tain that  of  all  the  differences  between  man  and  the  lower  animals, 
the  moral  sense  or  conscience  is  by  far  the  most  important.  This 
sense,  as  Mackintosh^  remarks,  "has  a  rightful  supremacy  over 
"every  other  principle  of  human  action;"  it  is  summed  up  in  that 
short  but  imperious  word  ought,  so  full  of  high  significance.  It 
is  the  most  noble  of  all  the  attributes  of  man,  leading  him  with- 
out a  moment's  hesitation  to  risk  his  life  for  that  of  a  fellow- 
creature;  or  after  due  deliberation,  impelled  simply  by  the  deep 
feeling  of  right  or  duty,  to  sacrifice  it  in  some  great  cause.  Im- 
manuel  Kant  exclaims,  "Duty!  Wondrous  thought,  that  workest 
"neither  by  fond  insinuation,  flattery,  nor  by  any  threat,  but 
"merely  by  holding  up  thy  naked  law  in  the  soul,  and  so  extorting 
"for  thyself  always  reverence,  if  not  always  obedience;  before 
"whom  all  appetites  are  dumb,  however  secretly  they  rebel; 
"whence  thy  original?"^ 

This  great  question  has  been  discussed  by  many  writers*  of 

1  See,  for  instance,  on  this  subject,  Quatrefages,  'Unite  de  I'Bspece 
Humaine,'  1861,  p.  21,  &c. 

2  'Dissertation  on  Ethical  Philosophy,'  1837,  p.  231,  &c. 

3  'Metaphysics  of  Ethics,'  translated  by  J.  W.  Semple,  Edinburgh, 
1836,  p.  136. 

*  Mr.  Bain  gives  a  list  ('Mental  and  Moral  Science,'  1868,  p.  543-725)  of 
twenty-six  British  authors  who  have  written  on  this  subject,  and 
whose  names  are  familiar  to  every  reader;  to  these,  Mr.  Bain's  own 


MORAL    SENSE.  95 

consummate  ability;  and  my  sole  excuse  for  touching  on  it,  is 
the  impossibility  of  here  passing  it  over;  and  because,  as  far  as  I 
know,  no  one  has  approached  it  exclusively  from  the  side  of 
natural  history.  The  investigation  possesses,  also,  some  inde- 
pendent interest,  as  an  attempt  to  see  how  far  the  study  of  the 
lower  animals  throws  light  on  one  of  the  highest  psychical  facul- 
ties of  man. 

The  following  proposition  seems  to  me  in  a  high  degree  prob- 
able— namely,  that  any  animal  whatever,  endowed  with  well- 
marked  social  instincts,^  the  parental  and  filial  affections  being 
here  included,  would  inevitably  acquire  a  moral  sense  or  con- 
science, as  soon  as  its  intellectual  powers  had  become  as  well, 
or  nearly  as  well  developed,  as  in  man.  For,  firstly,  the  social 
instincts  lead  an  animal  to  take  pleasure  in  the  society  of  its 
fellows,  to  feel  a  certain  amount  of  sympathy  with  them,  and  to 
perform  various  services  for  them.  The  services  may  be  of  a 
definite  and  evidently  instinctive  nature;  or  there  may  be  only 
a  wish  and  readiness,  as  with  most  of  the  higher  social  animals, 
to  aid  their  fellows  in  certain  general  ways.  But  these  feelings 
and  services  are  by  no  means  extended  to  all  the  individuals  of 
the  same  species,  only  to  those  of  the  same  association.  Secondly, 
as  soon  as  the  mental  faculties  had  become  highly  developed, 
images  of  all  past  actions  and  motives  would  be  incessantly 
passing  through  the  brain  of  each  individual;  and  that  feeling 
of  dissatisfaction,  or  even  misery,  which  invariably  results,  as  we 

name,  and  those  of  Mr.  Lecky,  Mr.  Shadworth  Hodgson,  Sir  J.  Lub- 
bock, and  others,  might  be  added. 

2  Sir  B.  Brodie,  after  observing  that  man  is  a  social  animal  ('Psy- 
chological Enquiries,'  1854,  p.  192),  asks  the  pregnant  question,  "ought 
"not  this  to  settle  the  disputed  question  as  to  the  existence  of  a  moral 
"sense?"  Similar  ideas  have  probably  occurred  to  many  persons,  as 
they  did  long  ago  to  Marcus  Aurelius.  Mr.  J.  S.  Mill  speaks,  in  his 
celebrated  work,  'Utilitarianism,'  (1864,  pp.  45,  46),  of  the  social  feelings 
as  a  "powerful  natural  sentiment,"  and  as  "the  natural  basis  of  senti- 
"ment  for  utilitarian  morality."  Again  he  says,  "Like  the  other  ac- 
"quired  capacities  above  referred  to,  the  moral  faculty,  if  not  a  part  of 
"our  nature,  is  a  natural  out-growth  from  it;  capable,  like  them,  in  a 
"certain  small  degree  of  springing  up  spontaneously."  But  in  oppo- 
sition to  all  this,  he  also  remarks,  "if,  as  is  my  own  belief,  the  moral 
"feelings  are  not  innate,  but  acquired,  they  are  not  for  that  reason  less 
"natural."  It  is  with  hesitation  that  I  venture  to  differ  at  all  from  so 
profound  a  thinker,  but  it  can  hardly  be  disputed  that  the  social  feel- 
ings are  instinctive  or  innate  in  the  lower  animals;  and  why  should 
they  not  be  so  in  man?  Mr.  Bain  (see,  for  instance,  'The  Emotions  and 
the  Will,'  1865,  p.  481)  and  others  believe  that  the  moral  sense  is  ac- 
quired by  each  individual  during  his  lifetime.  On  the  general  theory 
of  evolution  this  is  at  least  extremely  improbable.  The  ignoring  of  all 
transmitted  mental  qualities  will,  as  it  seems  to  me,  be  hereafter 
judged  as  a  most  serious  blemish  in  the  works  of  Mr.  Mill. 


96  THE  DESCENT  OF  MAN. 

shall  hereafter  see,  from  any  unsatisfied  instinct,  would  arise, 
as  often  as  it  was  perceived  that  the  enduring  and  alvsrays  present 
social  instinct  had  yielded  to  some  other  instinct,  at  the  time 
stronger,  but  neither  enduring  in  its  nature,  nor  leaving  behind 
it  a  very  vivid  impression.  It  is  clear  that  many  instinctive  de- 
sires, such  as  that  of  hunger,  are  in  their  nature  of  short  dura- 
tion; and  after  being  satisfied,  are  not  readily  or  vividly  recalled. 
Thirdly,  after  the  power  of  language  had  been  acquired,  and  the 
wishes  of  the  community  could  be  expressed,  the  common  opinion 
how  each  member  ought  to  act  for  the  public  good,  would  natural- 
ly become  in  a  paramount  degree  the  guide  to  action.  But  it 
should  be  borne  in  mind  that  however  great  weight  we  may  attrib- 
ute to  public  opinion,  our  regard  for  the  approbation  and  dis- 
approbation of  our  fellows  depends  on  sympathy,  which,  as  we 
shall  see,  forms  an  essential  part  of  the  social  instinct,  and  is 
indeed  its  foundation-stone.  Lastly,  habit  in  the  individual  would 
ultimately  play  a  very  important  part  in  guiding  the  conduct  of 
each  member;  for  the  social  instinct,  together  with  sympathy, 
is,  like  any  other  instinct,  greatly  strengthened  by  habit,  and  so 
consequently  would  be  obedience  to  the  wishes  and  judgment  of 
the  community.  These  several  subordinate  propositions  must  now 
be  discussed,  and  some  of  them  at  considerable  length. 

It  may  be  well  first  to  premise  that  I  do  not  wish  to  maintain 
that  any  strictly  social  animal,  if  its  intellectual  faculties  were 
to  become  as  active  and  as  highly  developed  as  in  man,  v/ould 
acquire  exactly  the  same  moral  sense  as  ours.  In  the  same  man- 
ner as  various  animals  have  some  sense  of  beauty,  though  they 
admire  widely  different  objects,  so  they  might  have  a  sense  of 
right  and  wrong,  though  led  by  it  to  follow  widely  different  lines 
of  conduct.  If,  for  instance,  to  take  an  extreme  case,  men  were 
reared  under  precisely  the  same  conditions  as  hive-bees,  there 
can  hardly  be  a  doubt  that  our  unmarried  females  would,  like 
the  worker-bees,  think  it  a  sacred  duty  to  kill  their  brothers,  and 
mothers  would  strive  to  kill  their  fertile  daughters;  and  no  one 
would  think  of  interfering."    Nevertheless,  the  bee,  or  any  other 


«  Mr.  H.  Sidg-wick  remarks  in  an  able  discussion  on  this  subject  (the 
'Academy,'  June  15th,  1872,  p.  231),  "a  superior  bee,  we  may  feel  sure, 
"would  aspire  to  a  milder  solution  of  the  population  question."  Judg- 
ing, however,  from  the  habits  of  many  or  most  savages,  man  solves  the 
problem  by  female  infanticide,  polyandry  and  promiscuous  inter- 
course; therefore  it  may  well  be  doubted  whether  it  would  be  by  a 
milder  method.  Miss  Cobbe,  in  commenting  ('Darwinism  in  Morals,' 
'Theological  Review,'  April,  1872,  p.  188-191)  on  the  same  illustration, 
says,  the  principles  of  social  duty  would  be  thus  reversed;  and  by  this, 
I  presume,  she  means  that  the  fulfillment  of  a  social  duty  would  tend 
to  the  injury  of  individuals;  but  she  overlooks  the  fact,  which  she 
would  doubtless  admit,   that  the  instincts  of  the  bee  have  been  ac- 


MORAL   SENSE.  97 

social  animal,  would  gain  in  our  supposed  case,  as  it  appears  to 
me,  some  feeling  of  right  or  wrong,  or  a  conscience.  For  each  in- 
dividual would  have  an  inward  sense  of  possessing  certain 
stronger  or  more  enduring  instincts,  and  others  less  strong  or 
enduring;  so  that  there  would  often  be  a  struggle  as  to  which 
impulse  should  be  followed;  and  satisfaction,  dissatisfaction,  or 
even  misery  would  be  felt,  as  past  impressions  were  compared 
during  their  incessant  passage  through  the  mind.  In  this  case 
an  inward  monitor  would  tell  the  animal  that  it  would  have  been 
better  to  have  followed  the  one  impulse  rather  than  the  other. 
The  one  course  ought  to  have  been  followed,  and  the  other  ought 
not;  the  one  would  have  been  right  and  the  other  wrong;  but  to 
these  terms  I  shall  recur. 

Sociability. — Animals  of  many  kinds  are  social;  we  find  even 
distinct  species  living  together;  for  example,  some  American 
monkeys;  and  united  flocks  of  rooks,  jackdaws,  and  starlings. 
Man  shows  the  same  feeling  in  his  strong  love  for  the  dog,  which 
the  dog  returns  with  interest.  Every  one  must  have  noticed  how 
miserable  horses,  dogs,  sheep,  &c.,  are  when  separated  from 
their  companions,  and  what  strong  mutual  affection  the  two 
former  kinds,  at  least,  show  on  their  reunion.  It  is  curious  to 
speculate  on  the  feelings  of  a  dog,  who  will  rest  peacefully  for 
hours  in  a  room  with  his  master  or  any  of  the  family,  without 
the  least  notice  being  taken  of  him;  but  if  left  for  a  short  time 
by  himself,  barks  or  howls  dismally.  We  will  confine  our  atten- 
tion to  the  higher  social  animals;  and  pass  over  insects,  although 
some  of  these  are  social,  and  aid  one  another  in  many  important  .;jf 

ways.  The  most  common  mutual  service  in  the  higher  animals 
is  to  warn  one  another  of  danger  by  means  of  the  united  senses  1 
of  all.  Every  sportsman  knows,  as  Dr.  Jaeger  remarks,"  how  diffi- 
cult it  is  to  approach  animals  in  a  herd  or  troop.  Wild  horses 
and  cattle  do  not,  I  believe,  make  any  danger-signal;  but  the 
attitude  of  any  one  of  them  who  first  discovers  an  enemy,  warns 
the  others.  Rabbits  stamp  loudly  on  the  ground  with  their  hind- 
feet  as  a  signal:  sheep  and  chamois  do  the  same  with  their  fore- 
feet, uttering  likewise  a  whistle.  Many  birds,  and  some  mam- 
mals, post  sentinels,  which  in  the  case  of  seals  are  said^  generally 
to  be  the  females.  The  leader  of  a  troop  of  monkeys  acts  as  the  sen- 
quired  for  the  good  of  the  community.  She  goes  so  far  as  to  say  that 
if  the  theory  of  ethics  advocated  in  this  chapter  were  ever  generally 
accepted,  "I  cannot  but  believe  that  in  the  hour  of  their  triumph  would 
"be  sounded  the  knell  of  the  virtue  of  mankind!"  It  is  to  be  hoped 
that  the  belief  in  the  permanence  of  virtue  on  this  earth  is  not  held  by 
many  persons  on  so  weak  a  tenure. 

■7  'Die  Darwin' sche  Theorie,'  s.  101. 

8  Mr.  R.  Brown  in  'Froc.  Zoolog.  Soc'  1868,  p.  4(». 


98  THE   DESCENT  OF  MAN. 

tinel,  and  utters  cries  expressive  both  of  danger  and  of  safety." 
Social  animals  perform  many  little  services  for  each  other:  horses 
nibble,  and  cows  lick  each  other,  on  any  spot  which  itches:  monk- 
eys search  each  other  for  external  parasites;  and  Brehm  states 
that  after  a  troop  of  the  Cercopithecus  griseo-viridis  has  rushed 
through  a  thorny  brake,  each  monkey  stretches  itself  on  a  branch, 
and  another  monkey  sitting  by,  "conscientiously"  examines  its 
fur,  and  extracts  every  thorn  or  burr. 

Animals  also  render  more  important  services  to  one  another: 
thus  wolves  and  some  other  beasts  of  prey  hunt  in  packs,  and 
aid  one  another  in  attacking  their  victims.  Pelicans  fish  in  con- 
cert. The  Hama'dryas  baboons  turn  over  stones  to  find  insects, 
&c.;  and  when  they  come  to  a  large  one,  as  many  as  can  stand 
round,  turn  it  over  together  and  share  the  booty.  Social  animals 
mutually  defend  each  other.  Bull  bisons  in  N.  America,  when 
there  is  danger,  drive  the  cows  and  calves  into  the  middle  of  the 
herd,  whilst  they  defend  the  outside.  I  shall  also  in  a  future  chap- 
ter give  an  account  of  two  young  wild  bulls  at  Chillingham  at- 
tacking an  old  one  in  concert,  and  of  two  stallions  together  trying 
to  drive  av/ay  a  third  stallion  from  a  troop  of  mares.  In  Abyssinia, 
Brehm  encountered  a  great  troop  of  baboons,  who  were  crossing 
a  valley:  some  had  already  ascended  the  opposite  mountain,  and 
some  were  still  in  the  valley:  tlie  latter  were  attacked  by  the 
dogs,  but  the  old  males  immediately  hurried  down  from  the  rocks, 
and  with  mouths  widely  opened  roared  so  fearfully,  that  the  dogs 
quickly  drew  back.  They  were  again  encouraged  to  the  attack; 
but  by  this  time  all  the  baboons  had  reascended  the  heights,  ex- 
cepting a  young  one,  about  six  months  old,  who,  loudly  calling 
for  aid,  climbed  on  a  block  of  rock,  and  was  surrounded.  Now 
one  of  the  largest  males,  a  true  hero,  came  down  again  from  the 
mountain,  slowly  went  to  the  young  one,  coaxed  him,  and  trium- 
phantly led  him  away — the  dogs  being  too  much  astonished  to 
make  an  attack.  I  cannot  resist  giving  another  scene  which  was 
witnessed  by  this  same  naturalist;  an  eagle  seized  a  young  Cerco- 
pithecus, which,  by  clinging  to  a  branch,  was  not  at  once  carried 
off;  it  cried  loudly  for  assistance,  upon  which  the  other  members 
of  the  troop,  with  much  uproar,  rushed  to  the  rescue,  surrounded 
the  eagle,  and  pulled  out  so  many  feathers,  that  he  no  longer 
thought  of  his  prey,  but  only  how  to  escape.    This  eagle,  as  Brehm 

8  Brehm,  'Thierleben,'  B.  i.  1864,  s.  52,  79.  For  the  case  of  the  mon- 
keys extracting  thorns  from  each  other,  see  s.  54.  With  respect  to  the 
Hamadryas  turning  over  stones,  the  fact  is  given  (s,  76)  on  the  evi- 
dence of  Alvarez,  whose  observations  Brehm  thinks  quite  trustworthy. 
For  the  cases  of  the  old  male  baboons  attacking  the  dogs,  see  s.  79; 
and  with  respect  to  the  eagle,  s.  56. 


MORAL.  SENSE.  99 

remarks,  assuredly  would  never  again  attack  a  single  monkey  of 
a  troop.^" 

It  is  certain  that  associated  animals  have  a  feeling  of  love  for 
each  other,  which  is  not  felt  by  non-social  adult  animals.  How 
far  in  most  cases  they  actually  sympathize  in  the  pains  and 
pleasures  of  others,  is  more  doubtful,  especially  with  respect  to 
pleasures.  Mr.  Buxton,  however,  who  had  excellent  means  of 
observation,^^  states  that  his  macaws,  which  lived  free  in  Norfolk, 
took  "an  extravagant  interest"  in  a  pair  with  a  nest;  and  when- 
ever the  female  left  it,  she  was  surrounded  by  a  troop  "scream- 
"ing  horrible  acclamations  in  her  honor."  It  is  often  difficult 
to  judge  whether  animals  have  any  feeling  for  the  sufferings  of 
others  of  their  kind.  Who  can  say  what  cows  feel,  when  they 
surround  and  stare  intently  on  a  dying  or  dead  companion;  ap- 
parently, however,  as  Houzeau  remarks,  they  feel  no  pity.  That 
animals  sometimes  are  far  from  feeling  any  sympathy  is  too 
certain;  for  they  will  expel  a  wounded  animal  from  the  herd,  or 
gore  or  worry  it  to  death.  This  is  almost  the  blackest  fact  in 
natural  history,  unless,  indeed,  the  explanation  which  has  been 
suggested  is  true,  that  their  instinct  or  reason  leads  them  to  expel 
an  injured  companion,  lest  beasts  of  prey,  including  man, 
should  be  tempted  to  follow  the  troop.  In  this  case  their  con- 
duct is  not  much  worse  than  that  of  the  North  American  Indians, 
who  leave  their  feeble  comrades  to  perish  on  the  plains:  or  the 
Fijians,  who,  when  their  parents  get  old,  or  fall  ill,  bury  them 
alive." 

Many  animals,  however,  certainly  sympathize  with  each  other's 
distress  or  danger.  This  is  the  case  even  with  birds.  Capt.  Stans- 
Ijui-yis  found  on  a  salt  lake  in  Utah  an  old  and  completely  blind 
pelican,  which  was  very  fat,  and  must  have  been  well  fed  for  a 
long  time  by  his  companions.  Mr.  Blyth,  as  he  informs  me,  saw 
Indian  crows  feeding  two  or  three  of  their  companions  which  were 
blind;    and  I  have  heard  of  an  analogous  case  with  the  domestic 

10  Mr.  Belt  gives  the  case  of  a  spider-monkey  (Ateles)  in  Nicara- 
gua which  was  heard  screaming  for  nearly  two  hours  in  the  forest, 
and  was  found  with  an  eagle  perched  close  by  it.  The  bird  apparently 
feared  to  attack  as  long  as  it  remained  face  to  face;  and  Mr.  Belt  be- 
lieves, from  what  he  has  seen  of  the  habits  of  these  monkeys,  that  they 
protect  themselves  from  eagles  by  keeping  two  or  three  together.  'The 
Naturalist  in  Nicaragua,'  1874,  p.  118. 

n  'Annals  and  Mag.  of  Nat.  Hist.,*  November,  1868,  p.  382. 

12  Sir  J.  Lubbock,  'Prehistoric  Times,'  2nd  edit.  p.  446. 

13  As  quoted  by  Mr.  L.  H.  Morgan,  'The  American  Beaver,*  1868,  p. 
272.  Capt.  Stansbury  also  gives  an  interesting  account  of  the  manner 
in  which  a  very  young  pelican,  carried  away  by  a  strong  stream,  was 
guided  and  encouraged  in  its  attempts  to  reach  the  shore  by  half  a 
dozen  old  birds. 


100  THE  DESCENT  OF  MAN. 

cock.  We  may,  if  we  choose,  call  these  actions  instinctive;  but 
such  cases  are  much  too  rare  for  the  development  of  any  special 
instinct/*  I  have  myself  seen  a  dog,  who  never  passed  a  cat  who 
lay  sick  in  a  basket,  and  was  a  great  friend  of  his,  without  giving 
her  a  few  licks  with  his  tongue,  the  surest  sign  of  kind  feeling  in 
a  dog. 

It  must  be  called  sympathy  that  leads  a  courageous  dog  to 
fly  at  any  one  who  strikes  his  master,  as  he  certainly  will.  I 
saw  a  person  pretending  to  beat  a  lady,  who  had  a  very  timid 
little  dog  on  her  lap,  and  the  trial  had  never  been  made  before; 
the  little  creature  instantly  jumped  away,  but  after  the  pretended 
beating  was  over,  it  was  really  pathetic  to  see  how  per&everingly 
he  tried  to  lick  his  mistress's  face,  and  comfort  her.  Brehm^* 
states  that  when  a  baboon  in  confinement  was  pursued  to  be 
punished,  the  others  tried  to  protect  him.  It  must  have  been 
sympathy  in  the  cases  above  given  which  led  the  baboons  and 
Cercopitheci  to  defend  their  young  comrades  from  the  dogs  and 
the  eagle.  I  will  give  only  one  other  instance  of  sympathetic 
and  heroic  conduct,  in  the  case  of  a  little  American  monkey. 
Several  years  ago  a  keeper  at  the  Zoological  Gardens  showed  me 
some  deep  and  scarcely  healed  wounds  on  the  nape  of  his  own 
neck,  inflicted  on  him,  whilst  kneeling  on  the  floor,  by  a  fierce 
baboon.  The  little  American  monkey,  who  was  a  warm  friend  of 
this  keeper,  lived  in  the  same  large  compartment,  and  was  dread- 
fully afraid  of  the  great  baboon.  Nevertheless,  as  soon  as  he  saw 
his  friend  in  peril,  he  rushed  to  the  rescue,  and  by  screams  and 
bites  so  distracted  the  baboon  that  the  man  was  able  to  escape, 
after,  as  the  surgeon  thought,  running  great  risk  of  his  life. 

Besides  love  and  sympathy,  animals  exhibit  other  qualities 
connected  with  social  instincts,  which  in  us  would  be  called  moral ; 
and  I  agree  with  Agassiz^^  that  dogs  possess  something  very  like 
a  conscience. 

Dogs  possess  some  power  of  self-command,  and  this  does  not 
appear  to  be  wholly  the  result  of  fear.  As  Braubach"  remarks, 
they  will  refrain  from  stealing  food  in  the  absence  of  their  mas- 
ter. They  have  long  been  accepted  as  the  very  type  of  fidelity 
and  obedience.  But  the  elephant  is  likewise  very  faithful  to  his 
driver  or  keeper,  and  probably  considers  him  as  the  leader  of 
the  herd.  Dr.  Hooker  informs  me  that  an  elephant,  which  he  was 
riding  in  India,  became  so  deeply  bogged  that  he  remained  stuck 
fast  until  the  next  day,  when  he  was  extricated  by  men  with 
ropes.    Under  such  circumstances  elephants  will  seize  with  their 

1*  As  Mr.  Bain  states,  "effective  aid  to  a  sufferer  springs  fronu.  sym- 
"pathy  proper;"  'Mental  and  Moral  Science,'  1868,  p.  245. 
15  'Thierleben,  B.  i.  s.  85. 
18  'De  I'Espece  et  de  la  Classe,'  1869,  p.  97. 
1^  'Die  Darwin' schfe  Art-Dehre,'  1869,  s.  54. 


MORAL  SENSE.  101 

trunks  any  object,  dead  or  alive,  to  place  under  their  knees,  to 
prevent  their  sinking  deeper  in  the  mud;  and  the  driver  was 
dreadfully  afraid  lest  the  animal  should  have  seized  Dr.  Hooker 
and  crushed  him  to  death.  But  the  driver  himself,  as  Dr.  Hooker 
was  assured,  ran  no  risk.  This  forbearance  under  an  emergency 
so  dreadful  for  a  heavy  animal,  is  a  wonderful  proof  of  noble 
fidelity.^^ 

All  animals  living  in  a  body,  which  defend  themselves  or  attack 
their  enemies  in  concert,  must  indeed  be  in  some  degree  faithful 
to  one  another;  and  those  that  follow  a  leader  must  be  in  some 
degree  obedient.  When  the  baboons  in  Abyssinia^^  plunder  a 
garden,  they  silently  follow  their  leader;  and  if  an  imprudent 
young  animal  makes  a  noise,  he  receives  a  slap  from  the  others 
to  teach  him  silence  and  obedience.  Mr.  Galton,  who  has  had 
excellent  opportunities  for  observing  the  half-wild  cattle  in  S. 
Africa,  says,-'^  that  they  cannot  endure  even  a  momentary  separa- 
tion from  the  herd.  They  are  essentially  slavish,  and  accept  the 
common  determination,  seeking  no  better  lot  than  to  be  led  by 
any  one  ox  who  has  enough  self-reliance  to  accept  the  position. 
The  men  who  break  in  these  animals  for  harness,  watch  assidu- 
ously for  those  who,  by  grazing  apart,  show  a  self-reliant  dis- 
position, and  these  they  train  as  fore-oxen.  Mr.  Galton  adds 
that  such  animals  are  rare  and  valuable;  and  if  many  were  born 
they  would  soon  be  eliminated,  as  lions  are  always  on  the  look- 
out for  the  individuals  wnich  wander  from  the  herd. 

With  respect  to  tne  impulse  which  leads  certain  animals  to 
associate  together,  and  to  aid  one  another  in  many  ways,  we  may 
infer  that  in  most  cases  they  are  impelled  by  the  same  sense  of 
satisfaction  or  pleasure  which  they  experience  in  performing 
other  instinctive  actions;  or  by  the  same  sense  of  dissatisfaction 
as  when  other  instinctive  actions  are  checked.  We  see  this  in 
innumerable  instances,  and  it  is  illustrated  in  a  striking  manner 
by  the  acquired  instincts  of  our  domesticated  animals;  thus  a 
young  shepherd-dog  delights  in  driving  and  running  round  a 
flock  of  sheep,  but  not  in  worrying  them;  a  young  fox-hound  de- 
lights in  hunting  a  fox,  whilst  some  other  kinds  of  dogs,  as  I  have 
witnessed,  utterly  disregard  foxes.  What  a  strong  feeling  of 
inward  satisfaction  must  impel  a  bird,  so  full  of  activity,  to  brood 
day  after  day  over  her  eggs.  Migratory  birds  are  quite  miserable 
if  stopped  from  migrating;  perhaps  they  enjoy  starting  on  their 
long  flight;  but  it  is  hard  to  believe  that  the  poor  pinioned  goose, 
described  by  Audubon,  which  started  on  foot  at  the  proper  time 
for  its  journey  of  probably  more  than  a  thousand  miles,  could 

^^  See  also  Hooker's  'Himalayan  Journals,'  vol.  ii.,  1854,  p.  333. 
1®  Brehm,  'Thierleben,'  B.  i.  s.  76. 

^  See  his  extremely  interesting-  paper  on  'Gregarlousness  in  Cattle, 
and  in  man,*  'Macmilian's  Meg."  Feb.  1871,  p.  353. 
8 


102  THE  DESCENT  OF  MAN. 

have  felt  any  joy  in  doing  so.  Some  instincts  are  determine<< 
solely  by  painful  feelings,  as  by  fear,  which  leads  to  self-preserva- 
tion, and  is  in  some  cases  directed  towards  special  enemies.  No 
one,  I  presume,  can  analyze  the  sensations  of  pleasure  or  pain. 
In  many  instances,  however,  it  is  probable  that  instincts  are  per- 
sistently followed  from  the  mere  force  of  inheritance,  without  the 
stimulus  of  either  pleasure  or  pain.  A  young  pointer,  when  it 
first  scents  game,  apparently  cannot  help  pointing.  A  squirrel 
in  a  cage  who  pats  the  nuts  which  it  cannot  eat,  as  if  to  bury  them 
in  the  ground,  can  hardly  be  thought  to  act  thus,  either  from 
pleasure  or  pain.  Hence  the  common  assumption  that  men  must 
be  impelled  to  every  action  by  experiencing  some  pleasure  or  pain 
may  be  erroneous.  Although  a  habit  may  be  blindly  and  im- 
plicitly followed,  independently  of  any  pleasure  or  pain  felt  at  the 
moment,  yet  if  it  be  forcibly  and  abruptly  checked,  a  vague  sense 
of  dissatisfaction  is  generally  experienced. 

It  has  often  been  assumed  that  animals  were  in  the  first  place 
rendered  social,  and  that  they  feel  as  a  consequence  uncomfortable 
when  separated  from  each  other,  and  comfortable  whilst  together; 
but  it  is  a  more  probable  view  that  these  sensations  were  first 
developed,  in  order  that  those  animals  which  would  profit  by  liv- 
ing in  society,  should  be  induced  to  live  together,  in  the  same 
manner  as  the  sense  of  hunger  and  the  pleasure  of  eating  v/ere. 
no  doubt,  first  acquired  in  order  to  induce  animals  to  eat.    The 
tfeeling  of  pleasure  from  society  is  probably  an  extension  of  the 
Iparental  or  filial  affections,  since  the  social  instinct  seems  to  be 
developed  by  the  young  remaining  for  a  long  time  with  their 
parents;    and  this  extension  may  be  attributed  in  part  to  habit, 
but  chiefly  to  natural  selection.    With  those  animals  which  were 
benefited  by  living  in  close  association,  the  individuals  which 
took  the  greatest  pleasure  in  society  would  best  escape  various 
dangers;    whilst  those  that  cared  least  for  their  comrades,  and 
lived  solitary,  would  perish  in  greater  numbers.   With  respect  to 
the  origin  of  the  parental  and  filial  affections,  which  apparently 
lie  at  the  base  of  the  social  instincts,  we  know  not  the  steps  by 
which  they  have  been  gained;    but  we  may  infer  that  it  has  been 
to  a  large  extent  through  natural  selection.     So  it  has  almost 
certainly  been  with  the  unusual  and  opposite  feeling  of  hatred 
between  the  nearest  relations,  as  with  the  worker-bees  which  kill 
their  brother-drones,  and  with  the  queen-bees  which  kill  their 
daughter-queens;    the  desire  to  destroy  their  nearest  relations 
having  been  in  this  case  of  service  to  the  community.    Parental 
affection,  or  some  feeling  which  replaces  it,  has  been  developed 
in  certain  animals  extremely  low  in  the  scale,  for  example,  in 
star-fishes  and  spiders.     It  is  also  occasionally  present  in  a  few 
members  alone  in  a  whole  group  of  animals,  as  in  the  genus  For- 
ficula,  or  earwigs. 


MORAL  SENSE.  103 

The  all-important  emotion  of  sympathy  is  distinct  from  that 
of  love.  A  mother  may  passionately  love  her  sleeping  and  passive 
infant,  but  she  can  hardly  at  such  times  be  said  to  feel  sympathy 
for  it.  The  love  of  a  man  for  his  dog  is  distinct  from  sympathy, 
and  so  is  that  of  a  dog  for  his  master.  Adam  Smith  formerly 
argued,  as  has  Mr.  Bain  recently,  that  the  basis  of  sympathy  lies 
in  our  strong  retentiveness  of  former  states  of  pain  or  pleasure. 
Hence,  "the  sight  of  another  person  enduring  hunger,  cold,  fa- 
"tigue,  revives  in  us  some  recollection  of  these  states,  which  are 
"painful  even  in  idea."  We  are  thus  impelled  to  relieve  the  suf- 
ferings of  another,  in  order  that  our  own  painful  feelings  may  be 
at  the  same  time  relieved.  In  like  manner  we  are  led  to  partici- 
pate in  the  pleasures  of  others.^^  But  I  cannot  see  how  this  view 
explains  the  fact  that  sympathy  is  excited,  in  an  immeasurably 
stronger  degree,  by  a  beloved,  than  by  an  indifferent  person.  The 
mere  sight  of  suffering,  independently  of  love,  would  suffice  to  call 
up  in  us  vivid  recollections  and  associations.  The  explanation 
may  lie  in  the  fact  that,  with  all  animals,  sympathy  is  directed 
solely  towards  the  members  of  the  same  community,  and  therefore 
towards  known,  and  more  or  less  beloved  members,  but  not  to  all 
the  individuals  of  the  same  species.  This  fact  is  not  more  sur- 
prising than  that  the  fears  of  many  animals  should  be  directed 
against  special  enemies.  Species  which  are  not  social,  such  as 
lions  and  tigers,  no  doubt  feel  sympathy  for  the  suffering  of  their 
own  young,  but  not  for  that  of  any  other  animal.  With  mankind, 
selfishness,  experience,  and  imitation,  probably  add,  as  Mr.  Bain 
has  shown,  to  the  power  of  sympathy;  for  we  are  led  by  the  hope 
of  receiving  good  in  return  to  perform  acts  of  sympathetic  kind- 
ness to  others;  and  sympathy  is  much  strengthened  by  habit.  In 
however  complex  a  manner  this  feeling  may  have  originated,  as 
it  is  one  of  high  importance  to  all  those  animals  which  aid  and 
defend  one  another,  it  will  have  been  increased  through  natural 
selection;  for  those  communities,  which  included  the  greatest 
number  of  the  most  sympathetic  members,  would  flourish  best, 
and  rear  the  greatest  number  of  offspring. 

It  is,  however,  impossible  to  decide  in  many  cases  whether  cer- 
tain social  instincts  have  been  acquired  through  natural  selection, 
or  are  the  indirect  result  of  other  instincts  and  faculties,  such  as 

21  See  the  first  and  striking  chapter  in  Adam  Smith's  'Theory  of 
Moral  Sentiments.'  Also  Mr.  Bain's  'Mental  and  Moral  Science,'  1868,  p. 
244,  and  275-282.  Mr.  Bain  states,  that  "sympathy  is,  indirectly,  a 
"source  of  pleasure  to  the  sympathizer;"  and  he  accounts  for  this 
through  reciprocity.  He  remarks  that  "the  person  benefited,  or  others 
"in  his  stead,  may  make  up  by  sympathy  and  good  offices  returned,  for 
"all  the  sacrifice."  But  if,  as  appears  to  be  the  case,  sympathy  is  strictly 
an  instinct,  its  exercise  would  give  direct  pleasure  in  the  same  manner, 
as  the  exercise,  as  before  remarked,  of  almost  every  other  instinct. 


104  THE  DESCENT  OF  MAN. 

sympathy,  reason,  experience,  and  a  tendency  to  imitation;  or 
again,  whether  they  are  simply  the  result  of  long-continued  habit. 
So  remarkable  an  instinct  as  the  placing  sentinels  to  warn  the 
community  of  danger,  can  hardly  have  been  the  indirect  result 
of  any  of  these  faculties;  it  must,  therefore,  have  been  directly 
acquired.  On  the  other  hand,  the  habit  followed  by  the  males  of 
some  social  animals  of  defending  the  community,  and  of  attacking 
their  enemies  or  their  prey  in  concert,  may  perhaps  have  orig- 
inated from  mutual  sympathy;  but  courage,  and  in  most  cases 
strength,  must  have  been  previously  acquired,  probably  through 
natural  selection. 

Of  the  various  instincts  and  habits,  some  are  much  stronger 
than  others;  that  is,  some  either  give  more  pleasure  in  their  per- 
formance, and  more  distress  in  their  prevention,  than  others,  or, 
which  is  probably  quite  as  important,  they  are,  through  inherit- 
ance, more  persistently  followed,  without  exciting  any  special  feel- 
ing of  pleasure  or  pain.  We  are  ourselves  conscious  that  some  hab- 
-its  are  much  more  difficult  to  cure  or  change  than  others.  Hence  a 
struggle  may  often  be  observed  in  animals  between  different  in- 
stincts, or  between  an  instinct  and  some  habitual  disposition;  as 
when  a  dog  rushes  after  a  hare,  is  rebuked,  pauses,  hesitates,  pur- 
sues again,  or  returns  ashamed  to  his  master;  or  as  between  the 
love  of  a  female  dog  for  her  young  puppies  and  for  her  master, 
— for  she  may  be  seen  to  slink  away  to  them,  as  if  half  ashamed 
of  not  accompanying  her  master.  But  the  most  curious  instance 
known  to  me  of  one  instinct  getting  the  better  of  another,  is  the 
migratory  instinct  conquering  the  maternal  instinct.  The  former 
is  wonderfully  strong;  a  confined  bird  will  at  the  proper  season 
beat  her  breast  against  the  wires  of  her  cage,  until  it  is  bare  and 
bloody.  It  causes  young  salmon  to  leap  out  of  the  fresh  water,  in 
which  they  could  continue  to  exist,  and  thus  unintentionally  to 
commit  suicide.  Every  one  knows  how  strong  the  maternal  in- 
stinct is,  leading  even  timid  birds  to  face  great  danger,  though 
with  hesitation,  and  in  opposition  to  the  instinct  of  self-preserva- 
tion. Nevertheless,  the  migratory  instinct  is  so  powerful,  that 
late  in  the  autumn  swallows,  house-martins,  and  swifts  frequently 
desert  their  tender  young,  leaving  them  to  perish  miserably  in 
their  nests.-^ 

^2  This  fact,  the  Rev.  L.  Jenyns  states  (see  his  edition  of  'White's 
Nat.  Hist,  of  Selborne,'  1853,  p.  204)  was  first  recorded  by  the  illustrious 
Jenner,  in  'Phil.  Transact.'  1824,  and  has  since  fceen  confirmed  by  sev- 
eral observers,  especially  by  Mr.  Blackwall.  This  latter  careful  ob- 
server examined  late  in  the  autumn,  during  two  years,  thirty-six  nests; 
he  found  that  twelve  contained  young-  dead  birds,  five  contained  eggs 
on  the  point  of  being  hatched,  and  three  eggs  not  nearly  hatched. 
Many  birds  not  yet  old  enough  for  a  prolonged  flight,  are  likewise 
desierted  and  lieift  b'efiind.    Steb  Blaokwall,  'Researches  in  ZoolDgy,'  1834, 


MORAL   SENSE.  105 

We  can  perceive  that  an  instinctive  impulse,  if  it  be  in  any- 
way more  beneficial  to  a  species  than  some  other  or  opposed 
instinct,  would  be  rendered  the  more  potent  of  the  two  through 
natural  selection;  for  the  individuals  which  had  it  most  strongly 
developed  would  survive  in  larger  numbers.  Whether  this  is  the 
case  with  the  migratory  in  comparison  with  the  maternal  instinct, 
may  be  doubted.  The  great  persistence,  or  steady  action  of  the 
former  at  certain  seasons  of  the  year  during  the  whole  day,  may 
give  it  for  a  time  paramount  force. 

Man  a  social  animal.  — Every  one  will  admit  that  man  is  a 
social  being.  We  see  this  in  his  dislike  of  solitude,  and  in  his 
wish  for  society  beyond  that  of  his  own  family.  Solitary  con- 
finement is  one  of  the  severest  punishments  which  can  be  inflicted. 
Some  authors  suppose  that  man  primevally  lived  in  single  fami- 
lies; but  at  the  present  day,  though  single  families,  or  only  two 
or  three  together,  roam  the  solitudes  of  some  savage  lands,  they 
always,  as  far  as  I  can  discover,  hold  friendly  relations  with 
other  families  inhabiting  the  same  district.  Such  families  oc- 
casionally meet  in  council,  and  unite  for  their  common  de- 
fense. It  is  no  argument  against  savage  man  being  a  social  ani- 
mal, that  the  tribes  inhabiting  adjacent  districts  are  almost  al- 
ways at  war  with  each  other;  for  the  social  instincts  never  ex- 
tend to  all  the  individuals  of  the  same  species.  Judging  from  the 
analogy  of  the  majority  of  the  Quadrumana,  it  is  probable  that 
the  early  ape-like  progenitors  of  man  v/ere  likewise  social;  but 
this  is  not  of  much  importance  for  us.  Although  man,  as  he 
now  exists,  has  few  special  instincts,  having  lost  any  which  his 
early  progenitors  may  have  possessed,  this  is  no  reason  why  he 
should  not  have  retained  from  an  extremely  remote  period  some 
degree  of  instinctive  love  and  sympathy  for  his  fellows.  We 
are  indeed  all  conscious  that  we  do  possess  such  sympathetic 
feelings;^  but  our  consciousness  does  not  tell  us  whether  they 
are  instinctive,  having  originated  long  ago  in  the  same  manner 
as  with  the  lower  animals,  or  whether  they  have  been  acquired 
by  each  of  us  during  our  early  years.     As  man  is  a  social  animal, 

pp.  108,  118.  For  some  additional  evidence,  althoug-h  this  is  not  wanted, 
see  Leroy,  'Lettres  Pliil.'  1802,  p.  217.  For  Swifts,  Goulds,  'IntrodLiction 
to  the  Birds  of  Great  Britain,'  1823,  p.  5.  Similar  cases  have  been  ob- 
served in  Canada  by  Mr.  Adams;  'Pop.  Science  Review,'  July,  1873, 
p.  283. 

23  Hume  remarks  ('An  Enquiry  Concerning  the  Principles  of  Morals,' 
edit,  of  1751,  p.  132).  "There  seems  a  necessity  for  confessing  that  the 
"happiness  and  misery  of  others  are  not  spectacles  altogether  indiffer- 
"ent  to  us,  but  that  the  view  of  the  former  .  ,  .  communicates  a  secret 
"joy;  the  appearance  of  the  latter  .  .  .  throws  a  melancholy  damp 
"over  the  imagination." 


106  THE  DESCENT  OF   MAN. 

it  is  almost  certain  that  he  would  inherit  a  tendency  to  be  faith- 
ful to  his  comrades,  and  obedient  to  the  leader  of  his  tribe  for 
these  qualities  are  common  to  most  social  animals.  He  would 
consequently  possess  some  capacity  for  self-command.  He 
would  from  an  inherited  tendency  be  willing  to  defend,  in  con- 
cert with  others,  his  fellow-men;  and  would  be  ready  to  aid 
them  in  any  way,  which  did  not  too  greatly  interfere  with  his 
own  welfare  or  his  own  strong  desires. 

The  social  animals  which  stand  at  the  bottom  of  the  scale  are 
guided  almost  exclusively,  and  those  which  stand  higher  in  the 
scale  are  largely  guided,  by  special  instincts  in  the  aid  which 
they  give  to  the  members  of  the  same  community;  but  they  are 
likewise  in  part  impelled  bj;^  mutual  love  and  sympathy,  assisted 
apparently  by  some  amount  of  reason.  Although  man,  as  just 
remarked,  has  no  special  instincts  to  tell  him  how  to  aid  his 
fellow-men,  he  still  has  the  impulse,  and  with  his  improved 
intellectual  faculties  would  naturally  be  much  guided  in  this  re- 
spect by  reason  and  experience.  Instinctive  sympathy  would 
also  cause  him  to  value  highly  the  approbation  of  his  fellows; 
for,  as  Mr.  Bain  has  clearly  shown,-*  the  love  of  praise  and  the 
strong  feeling  of  glory,  and  the  still  stronger  horror  of  scorn 
and  infamy,  "are  due  to  the  workings  of  sympathy."  Conse- 
quently man  would  be  influenced  in  the  highest  degree  by  the 
wishes,  approbation,  and  blame  of  his  fellow-men,  as  expressed 
by  their  gestures  and  language.  Thus  the  social  instincts, 
which  must  have  been  acquired  by  man  in  a  very  rude  state, 
and  probably  even  by  his  early  ape-like  progenitors,  still  give 
the  impulse  to  some  of  his  best  actions;  but  his  actions  are  in  a 
higher  degree  determined  by  the  expressed  wishes  and  judgment 
of  his  fellow-men,  and  unfortunately  very  often  by  his  own 
strong  selfish  desires.  JBut  as  love,  sympathy  and  self-command 
become  strengthened  by  habit,  and  as  the  power  of  reasoning 
becomes  clearer,  so  that  man  can  value  justly  the  judgments  of 
his  fellows,  he  will  feel  himself  impelled,  apart  trom  any  transi- 
tory pleasure  or  pain,  to  certain  lines  of  conduct.  He  might 
then  declare — not  that  any  barbarian  or  uncultivated  man  could 
thus  think — I  am  the  supreme  judge  of  my  own  conduct,  and  in 
the  words  of  Kant,  I  will  not  in  my  own  person  violate  the 
dignity  of  humanity. 

The  more  enduring  Social  Instincts  conquer  the  less  persistent 
Instincts.  — We  have  not,  however,  as  yet  considered  the  main 
point,  on  which,  from  our  present  point  of  view,  the  whole 
question  of  the  moral  sense  turns.  Why  should  a  man  feel  that 
he  ought  to   obey   one   instinctive   desire  rather  than  another? 

24  'Mental  and  Moral  Science,'  1868,  p.  254. 


MORAL  SENSE.  107 

Why  is  he  bitterly  regretful,  if  he  has  yielded  to  a  strong  sense 
of  self-preservation,  and  has  not  risked  his  life  to  save  that  of  a 
fellow-creature?  or  why  does  he  regret  having  stolen  food  from 
hunger? 

It  is  evident  in  the  first  place,  that  with  mankind  the  instinc- 
tive impulses  have  different  degrees  of  strength;  a  savage  will 
risk  his  own  life  to  save  that  of  a  member  of  the  same  com- 
munity, but  will  be  wholly  indifferent  about  a  stranger:  a  young 
and  timid  mother  urged  by  the  maternal  instinct  will,  without 
a  moment's  hesitation,  run  the  greatest  danger  for  her  own  in- 
fant, but  not  for  a  mere  fellow-creature.  Nevertheless  many  a 
civilized  man,  or  even  boy,  who  never  before  risked  his  life  for 
another,  but  full  of  courage  and  sympathy,  has  disregarded 
the  instinct  of  self-preservation,  and  plunged  at  once  into  a 
torrent  to  save  a  drowning  man,  though  a  stranger.  In  this  case 
man  is  impelled  by  the  same  instinctive  motive,  which  made  the 
heroic  little  American  monkey,  formerly  described,  save  his 
keeper,  by  attacking  the  great  and  dreaded  baboon.  Such  ac- 
tions as  the  above  appear  to  be  the  simple  result  of  the  greater 
strength  of  the  social  or  maternal  instincts  than  that  of  any 
other  instinct  or  motive;  for  they  are  performed  too  instan- 
taneously for  reflection,  or  for  pleasure  or  pain  to  be  felt  at  the 
time;  though,  if  prevented  by  anj^  cause,  distress  or  even  misery 
might  be  felt.  In  a  timid  man,  on  the  other  hand,  the  instinct 
of  self-preservation  might  be  so  strong,  that  he  would  be  unable 
to  force  himself  to  run  any  such  risk,  perhaps  not  even  for  his 
own-child. 

I  am  aware  that  some  persons  maintain  that  actions  performed 
impulsively,  as  in  the  above  cases,  do  not  come  under  the  do- 
minion of  the  moral  sense,  and  cannot  be  called  moral.  They 
confine  this  term  to  actions  done  deliberately,  after  a  victory 
over  opposing  desires,  or  when  prompted  by  some  exalted  mo- 
tive. But  it  appears  scarcely  possible  to  draw  any  clear  line 
of  distinction  of  this  kind.-^  As  far  as  exalted  motives  are 
concerned,  many  instances  have  been  recorded  of  savages,  des- 
titute of  any  feeling  of  general  benevolence  towards  mankind, 
and  not  guided  by  any  religious  motive,  who  have  deliberately  . 
sacrificed    their   lives   as    prisoners,^**   rather   than   betray   their  ^; 

25 1  refer  here  to  the  distinction  between  what  has  been  called  ma- 
terial and  formal  morality.  I  am  glad  to  And  that  Prof.  Huxley 
('Critiques  and  Addresses,'  1873,  p.  287)  takes  th©  same  view  on  this 
subject  as  I  do.  Mr.  Leslie  Stephen  remarks  ('Essays  on  Preethinking- 
and  Plain  Speaking,'  1873,  p.  S3),  "the  metaphysical  distinction  be- 
"tween  material  and  formal  morality  is  as  irrelevant  as  other  such  dis- 
"tinctions." 

^  I  have  given  on©  such  case,  namely  of  three  Patagoniaa  Indians 


108  THE  DESCENT  OP  MAN. 

comrades;  and  surely  their  conduct  ought  to  be  considered  as 
moral.  As  far  as  deliberation,  and  the  victory  over  opposing 
motives  are  concerned,  animals  may  be  seen  doubting  between 
opposed  instincts,  in  rescuing  their  offspring  or  comrades  from 
danger;  yet  their  actions,  though  done  for  the  good  of  others, 
are  not  called  moral.  Moreover,  anything  performed  very  often 
by  us,  will  at  last  be  done  without  deliberation  or  hesitation, 
and  can  then  hardly  be  distinguished  from  an  instinct;  yet 
surely  no  one  will  pretend  that  such  an  action  ceases  to  be 
moral.  On  the  contrary,  we  all  feel  that  an  act  cannot  be 
considered  as  perfect,  or  as  performed  in  the  most  noble  manner, 
unless  it  be  done  impulsively,  without  deliberation  or  effort,  in 
the  same  manner  as  by  a  man  in  whom  the  requisite  qualities  are 
innate.  He  who  is  forced  to  overcome  his  fear  or  want  of  sym- 
pathy before  he  acts,  deserves,  however,  in  one  way  higher 
credit  than  the  man  whose  innate  disposition  leads  him  to  a 
good  act  without  effort.  As  we  cannot  distinguish  between 
motives,  we  rank  all  actions  of  a  certain  class  as  moral,  if  per- 
formed by  a  moral  being.  A  moral  being  is  one  who  is  capable 
of  comparing  his  past  and  future  actions  or  motives,  and  of 
approving  or  disapproving  of  them.  We  have  no  reason  to  sup- 
pose that  any  of  the  lower  animals  have  this  capacitj'-;  there- 
fore, when  a  Newfoundland  dog  drags  a  child  out  of  the  water, 
or  a  monkey  faces  danger  to  rescue  its  comrade,  or  takes  charge 
of  an  orphan  monkey,  we  do  not  call  its  conduct  moral.  But 
in  the  case  of  man,  who  alone  can  with  certainty  be  ranked 
as  a  moral  being,  actions  of  a  certain  class  are  called  moral, 
whether  performed  deliberately,  after  a  struggle  with  opposing 
motives,  or  impulsively  through  instinct,  or  from  the  effects  of 
slowly-gained  habit. 

But  to  return  to  our  more  immediate  subject.  Although  some 
instincts  are  more  powerful  than  others,  and  thus  lead  to  cor- 
responding actions,  yet  it  is  untenable,  that  in  man  the  social 
instincts  (including  the  love  of  praise  and  fear  of  blame)  possess 
greater  strength,  or  have,  through  long  habit,  acquired  greater 
strength  than  the  instincts  of  self-preservation,  hunger,  lust, 
vengeance,  &c.  Why  then  does  man  regret,  even  though  trying 
to  banish  such  regret,  that  he  has  followed  the  one  natural 
impulse  rather  than  the  other;  and  why  does  he  further  feel 
that  he  ought  to  regret  his  conduct?  Man  in  this  respect  differs 
profoundly  from  the  lower  animals.  Nevertheless  we  can  I  think, 
see  with  some  degree  of  clearness  the  reason  of  this  difference. 

Man,  from  the  activity  of  his  mental  faculties,  cannot  avoid 
reflection:     past    impressions    and    images  are  incessantly    and 

who  preferred  being  shot,  one  after  the  other,  to  betraying  the  plans 
Of  their  companions  in  war  ('Journai  of  Researches,'  1845,  p.  103), 


MORAL  SENSE.  109 

clearly  passing  through  his  mind.  Now  with  those  animals 
which  live  permanently  in  a  body,  the  social  instincts  are  ever 
present  and  persistent.  Such  animals  are  always  ready  to  utter 
the  danger-signal,  to  defend  the  community,  and  to  give  aid  to 
their  fellows  in  accordance  with  their  habits;  they  feel  at  all 
times,  without  the  stimulus  of  any  special  passion  or  desire, 
some  degree  of  love  and  sympathy  for  them;  they  are  unhappy 
if  long  separated  from  them,  and  always  happy  to  be  again  in 
their  company.  So  it  is  with  ourselves.  Even  when  we  are 
quite  alone,  how  often  do  we  think  with  pleasure  or  pain  of  what 
others  think  of  us, — of  their  imagined  approbation  or  disappro- 
bation; and  this  all  follows  from  sympathy,  a  fundamental 
element  of  the  social  instincts.  A  man  who  possessed  no  trace 
of  such  instincts  would  be  an  unnatural  monster.  On  the  other 
hand,  the  desire  to  satisfy  hunger,  or  any  passion  such  as  ven- 
geance, is  in  its  nature  temporary,  and  can  for  a  time  be  fully 
satisfied.  Nor  is  it  easy,  perhaps  hardly  possible,  to  call  up 
with  complete  vividness  the  feeling,  for  instance,  of  hunger;  nor 
indeed,  as  has  often  been  remarked,  of  any  suffering.  The  in- 
stinct of  self-preservation  is  not  felt  except  in  the  presence  of 
danger;  and  many  a  coward  has  thought  himself  brave  until  he 
has  met  his  enemy  face  to  face.  The  wish  for  another  man's 
property  is  perhaps  as  persistent  a  desire  as  any  that  can  be 
named;  but  even  in  this  case  the  satisfaction  of  actual  pos- 
session is  generally  a  weaker  feeling  than  the  desire:  many  a 
thief,  if  not  a  habitual  one,  after  success  has  v/ondered  why  he 
stole  some  article." 


27  Enmity  or  hatred  seems  also  to  be  a  highly  persistent  feeling,  per- 
haps more  so  than  any  other  that  can  be  named.  Envy  is  defined  as 
hatred  of  another  for  some  excellence  or  success;  and  Bacon  insists 
(Essay  ix.),  "Of  all  other  aifections  envy  is  the  most  importune  and 
"continual."  Dogs  are  very  apt  to  hate  both  strange  men  and  strange 
dogs,  especially  if  they  live  near  at  hand,  but  do  not  belong  to  the  same 
family,  tribe,  or  clan;  this  feeling  would  thus  seem  to  be  innate  and 
is  certainly  a  most  persistent  one.  It  seems  to  be  the  complement  and 
converse  of  the  true  social  instinct.  From  what  we  hear  of  savages, 
it  would  appear  that  something  of  the  same  kind  holds  good  with  them. 
If  this  be  so,  it  would  be  a  small  step  in  any  one  to  transfer  such 
feelings  to  any  member  of  the  same  tribe  if  he  had  done  him  an 
injury  and  had  become  his  enemy.  Nor  is  it  probable  that  the  prim- 
itive conscience  would  reproach  a  man  for  injuring  his  enemy:  rather 
it  would  reproach  him  if  he  had  not  revenged  himself.  To  do  good  in 
return  for  evil,  to  love  your  enemy,  is  a  height  of  morality  to  which 
it  may  be  doubted  whether  the  social  instincts  would,  by  themselves, 
have  ever  led  us.  It  is  necessary  that  these  instincts,  together  with 
sympathy,  should  have  been  highly  cultivated  and  extended  by  the  aid 
of  reason,  instruction,  and  the  love  or  fear  of  God  before  any  such 
golden  rule  would  ever  be  thought  of  and  obeyed. 


110  THE  DESCENT  OF  MAN. 

A  man  cannot  prevent  past  impressions  often  repassing 
through  his  mind;  he  will  thus  be  driven  to  make  a  comparison 
between  the  impressions  of  past  hunger,  vengeance  satisfied,  or 
danger  shunned  at  other  men's  cost,  with  the  almost  ever-present 
instinct  of  sympathy,  and  with  his  early  knowledge  of  what  others 
consider  as  praisev/orthy  or  blamable.  This  knowledge  can- 
not be  banished  from  his  mind,  and  from  instinctive  sympathy 
is  esteemed  of  great  moment.  He  will  then  feel  as  if  he  had 
been  balked  in  following  a  present  instinct  or  habit,  and  this 
with  all  animals  causes  dissatisfaction,  or  even  misery. 

The  above  case  of  the  swallow  affords  an  illustration,  though 
of  a  reversed  nature,  of  a  temporary  though  for  the  time  strongly 
persistent  instinct  conquering  another  instinct,  which  is  usually 
dominant  over  all  others.  At  the  proper  season  these  birds 
seem  all  day  long  to  be  impressed  with  the  desire  to  migrate; 
their  habits  change;  they  become  restless,  are  noisy,  and  con- 
gregate in  flocks.  "Whilst  the  mother-bird  is  feeding,  or  brood- 
ing over  her  nestlings,  the  maternal  instinct  is  probably  stronger 
than  the  migratory;  but  the  instinct  which  is  the  more  persist- 
ent gains  the  victory,  and  at  last,  at  a  moment  when  her  young 
ones  are  not  in  sight,  she  takes  flight  and  deserts  them.  When 
arrived  at  the  end  of  her  long  journey,  and  the  migratory  in- 
stinct has  ceased  to  act,  what  an  agony  of  remorse  the  bird 
would  feel,  if,  from  being  endowed  with  great  mental  activity, 
she  could  not  prevent  the  image  constantly  passing  through  her 
mind,  of  her  young  ones  perishing  in  the  bleak  north  from  cold 
and  hunger. 

At  the  moment  of  action,  man  will  no  doubt  be  apt  to  follow 
the  stronger  impulse;  and  though  this  may  occasionally  prompt 
him  to  the  noblest  deeds,  it  will  more  commonly  lead  him  to 
gratify  his  own  desires  at  the  expense  of  other  men.  But  after 
their  gratification,  when  past  and  weaker  impressions  are  judged 
by  the  ever-enduring  social  instinct,  and  by  his  deep  regard 
for  the  good  opinion  of  his  fellows,  retribution  will  surely  come. 
He  will  then  feel  remorse,  repentance,  regret,  or  shame;  this 
latter  feeling,  however,  relates  almost  exclusively  to  the  judg- 
ment of  others.  He  will  consequently  resolve  more  or  less  firmly 
to  act  differently  for  the  future;  and  this  is  conscience;  for 
conscience  looks  backwards,  and  serves  as  a  guide  for  the  future. 

The  nature  and  strength  of  the  feelings  which  we  call  regret, 
shame,  repentance  or  remorse,  depend  apparently  not  only  on 
the  strength  of  the  violated  instinct,  but  partly  on  the  strength 
of  the  temptation,  and  often  still  more  on  the  judgment  of  our 
fellows.  How  far  each  man  values  the  appreciation  of  others, 
depends  on  the  strength  of  his  innate  or  acquired  feeling  of 
sympathy;  and  on  his  own  capacity  for  reasoning  out  the  re- 
mote consequences  of  his  acts.     Another  element  is  most  im- 


MORAL   SENSE.  Ill 

portant,  although  not  necessary,  the  reverence  or  fear  of  the 
Gods,  or  Spirits  believed  in  by  each  man:   and  this  applies  es- 
pecially in  cases  of  remorse.    Several  critics  have  objected  that 
though  some  slight  regret  or  repentance  may  be  explained  by 
the  view  advocated  in  this  chapter,  it  is  impossible  thus  to  ac- 
count for  the  soul-shaking  feeling  of  remorse.     But  I  can  see 
little  force  in  this   objection.     My   critics   do   not  define   what 
they  mean  by  remorse,  and  I  can  find  no  definition  implying 
more  than   an   overwhelming   sense   of    repentance.      Remorse 
seems  to  bear  the  same  relation  to  repentance,  as  rage  does  to 
anger,  or  agony  to  pain.    It  is  far  from  strange  that  an  instinct 
so  strong  and  so  generally  admired,  as  maternal  love,  should,  if 
disobeyed,  lead  to  the  deepest  misery,  as  soon  as  the  impression 
of  the  past  cause  of  disobedience  is  weakened.     Even  when  an 
action  is  opposed  to  no  special  instinct,  merely  to  know  that  our 
friends  and  equals  despise  us  for  it  is  enough  to  cause  great 
misery.    Who  can  doubt  that  the  refusal  to  fight  a  duel  through 
fear  has  caused  many  men  an  agony  of  shame?    Many  a  Hindoo, 
it  is  said,  has  been  stirred  to  the  bottom  of  his  soul  by  having 
partaken  of  unclean  food.    Here  is  another  case  of  what  must,  I 
think,  be  called  remorse.     Dr.  Landor  acted  as  a  magistrate  in 
West  Australia,  and  relates,-*^  that  a  native  on  his  farm,  after 
losing  one  of  his  wives  from  disease,  came  and  said  that  "he  was 
"going  to  a  distant  tribe  to  spear  a  woman,  to  satisfy  his  sense  | 
"of  duty  to  his  wife.     I  told  him  that  if  he   did  so,   I  Y\rouldi 
"send  him  to  prison  for  life.     He  remained  about  the  farm  for*^ 
"some  months,   but  got  exceedingly  thin,  and  complained  that 
"he  could  not  rest  or  eat,  that  his  wife's  spirit  was  haunting 
"him,  because  he  had  not  taken  a  life  for  hers.     I  was  inex- 
"orable,  and  assured  him  that  nothing  should  save  him  if  he 
"did."    Nevertheless  the  man  disappeared  for  more  than  a  year, 
and  then  returned  in  high  condition;    and  his  other  wife  told 
Dr.   Landor  that  her  husband  had  taken  the  life  of  a  woman 
belonging  to  a  distant  tribe;    but  it  was  impossible  to  obtain 
legal  evidence  of  the  act.     The  breach  of  a  rule  held  sacred  by 
the  tribe,  will  thus,  as  it  seems,  give  rise  to  the  deepest  feelings, 
— and  this  quite  apart  from  the  social  instincts,  excepting  in  so 
far  as  the  rule  is  grounded  on  the  judgment  of  the  community. 
How  so  many  strange  superstitions  have  arisen  throughout  the 
world  we  know  not;  nor  can  we  tell  how  some  real  and  great 
crimes,  such  as  incest,  have  come  to  be  held  in  an  abhorrence 
(which  is  not  however  quite  universal)  by  the  lowest  savages.    It 
Is  even  doubtful  whether  in  some  tribes  incest  would  be  looked  on 
with  greater  horror,  than  would  the  marriage  of  a  man  with  a 
woman   bearing  the   same  name,   though   not  a   relation.     "To 


'Insanity  in  Relation  to  Law;'  Ontario,  United  States,  1871,  p.  14. 


112  THE  DESCENT  OF  MAN. 

"violate  this  law  is  a  crime  which  the  Australians  hold  in  the 
"greatest  abhorrence,  in  this  agreeing  exactly  with  certain 
"tribes  of  North  America.  When  the  question  is  put  in  either 
"district,  is  it  worse  to  kill  a  girl  of  a  foreign  tribe,  or  to  marry 
"a  girl  of  one's  own,  an  answer  just  opposite  to  ours  would  be 
"given  without  hesitation.""'  We  may,  therefore,  reject  the  be- 
lief, lately  insisted  on  by  some  writers,  that  the  abhorrence  of 
incest  is  due  to  our  possessing  a  special  God-implanted  con- 
science. On  the  whole  it  is  intelligible,  that  a  man  urged  by 
so  powerful  a  sentiment  as  remorse,  though  arising  as  above 
explained,  should  be  led  to  act  in  a  manner,  which  he  has  been 
taught  to  believe  serves  as  an  expiation,  such  as  delivering  him- 
self up  to  justice. 

Man  prompted  by  his  conscience,  will  through  long  habit  ac- 
quire such  perfect  self-command,  that  his  desires  and  passions 
will  at  last  yield  instantly  and  without  a  struggle  to  his  social 
sympathies  and  instincts,  including  his  feeling  for  the  judgment 
of  his  fellows.  The  still  hungry,  or  the  still  revengeful  man  will 
not  think  of  stealing  food,  or  of  wreaking  his  vengeance.  It  is 
possible,  or  as  we  shall  hereafter  see,  even  probable,  that  the 
habit  of  self-command  may,  like  other  habits,  be  inherited.  Thus 
at  last  man  comes  to  feel,  through  acquired  and  perhaps  inherited 
habit,  that  it  is  best  for  him  to  obey  his  more  persistent  impulses. 
The  imperious  word  ought  seems  merely  to  imply  the  conscious- 
ness of  the  existence  of  a  rule  of  conduct,  however  it  may  have 
originated.  Formerly  it  must  have  been  often  vehemently  urged 
that  an  insulted  gentleman  ought  to  fight  a  duel.  We  even  say 
that  a  pointer  ought  to  point,  and  a  retriever  to  retrieve  game.  If 
they  fail  to  do  so,  they  fail  in  their  duty  and  act  wrongly. 

If  any  desire  or  instinct  leading  to  an  action  opposed  to  the 
good  of  others  still  appears,  when  recalled  to  mind,  as  strong 
as,  or  stronger  than,  the  social  instinct,  a  man  will  feel  no  keen 
regret  at  having  followed  it;  but  he  will  be  conscious  that  if  his 
conduct  were  known  to  his  fellows,  it  would  meet  with  their 
disapprobation;  and  few  are  so  destitute  of  sympathy  as  not  to 
feel  discomfort  when  this  is  realized.  If  he  has  no  such  sym- 
pathy, and  if  his  desires  leading  to  bad  actions  are  at  the  time 
strong,  and  when  recalled  are  not  over-mastered  by  the  persistent 
social  instincts,  and  the  judgment  of  others,  then  he  is  essentially 
a  bad  man;^*^  and  the  sole  restraining  motive  left  is  the  fear  of 
punishment,  and  the  conviction  that  in  the  long  run  it  would  be 
best  for  his  own  selfish  interests  to  regard  the  good  of  others 
rather  than  his  own. 


20  E.  B.  Tylor  in  'Contemporary  Heview,'  April,  1873,  p.  707. 

30  Dr.  Prosper  Despine,  in  his  'Psychologie  Naturelle,'  1868  (torn,  i,  p. 
243;  torn.  ii.  p.  169)  gives  many  curious  cases  of  the  worst  criminals, 
who  apparently  have  been  entirely  destitute  of  conscience. 


MORAL  SENSE.  113 

It  is  obvious  ttiat  every  one  may  with  an  easy  conscience 
gratify  his  own  desires,  if  they  do  not  interfere  with  his  social 
instincts,  that  is  with  the  good  of  others;  but  in  order  to  be 
quite  free  from  self-reproach,  or  at  least  of  anxiety,  it  is  almost 
necessary  for  him  to  avoid  the  disapprobation,  whether  reason- 
able or  not,  of  his  fellow-men.  Nor  must  he  break  through  the 
fixed  habits  of  his  life,  especially''  if  these  are  supported  by  reason; 
for  if  he  does,  he  will  assuredly  feel  dissatisfaction.  He  must 
likewise  avoid  the  reprobation  of  the  one  God  or  gods  in  whom, 
according  to  his  knowledge  or  superstition,  he  may  believe;  but 
in  this  case  the  additional  fear  of  divine  punishment  often  super- 
venes. 

The  strictly  Social  VlHues  at  first  alone  regarded. — The  above 
view  of  the  origin  and  nature  of  the  moral  sense,  which  tells  us 
what  we  ought  to  do,  and  of  the  conscience  which  reproves  us  if 
we  disobey  it,  accords  well  with  what  we  see  of  the  early  and 
undeveloped  condition  of  this  faculty  in  mankind.  The  virtues 
which  must  be  practiced,  at  least  generally,  by  rude  men,  so 
that  they  may  associate  in  a  body,  are  those  which  are  still 
recognized  as  the  most  important.  But  they  are  practiced  almost 
exclusively  in  relation  to  the  men  of  the  same  tribe;  and  their 
opposites  are  not  regarded  as  crimes  in  relation  to  the  men  of 
other  tribes.  No  tribe  could  hold  together  if  murder,  robbery, 
treachery,  &c.,  were  common;  consequently  such  crimes  within 
the  limits  of  the  same  tribe  "are  branded  with  everlasting  in- 
"famy;"^^  but  excite  no  such  sentiment  beyond  these  limits.  A 
North-American  Indian  is  well  pleased  with  himself,  and  is  hon- 
ored by  others,  when  he  scalps  a  man  of  another  tribe;  and  a 
Dj^ak  cuts  off  the  head  of  an  unoffending  person,  and  dries  it  as 
a  trophy.  The  murder  of  infants  has  prevailed  on  the  largest 
scale  throughout  the  world,^  and  has  met  with  no  reproach;  but 
infanticide,  especially  of  females,  has  been  thought  to  be  good 
for  the  tribe,  or  at  least  not  injurious.  Suicide  during  former 
times  was  not  generally  considered  as  a  crime,^^  but  rather,  from 

31  See  an  able  article  in  the  'North  British  Review,'  1S67,  p.  395.  See 
also  Mr,  W.  Bagehot's  articles  on  the  Importance  of  Obedience  and 
Coherence  to  Primitive  Man,  in  the  'Fortnightly  Review,'  IS'oT,  p.  529, 
and  1S68,  p.  457,  &c. 

^  The  fullest  account  which  I  have  met  with  is  by  Dr.  Gerland,  in 
his  'Ueber  das  Aussterben  der  Naturvolker,'  1868;  but  I  shall  have  to 
recur  to  the  subject  of  infanticide  in  a  future  chapter. 

^  See  the  very  interesting  discussion  on  Suicide  in  Lecky's  'History 
of  European  Morals,'  vol.  i.  1869,  p.  223..  With  respect  to  savages,  Mr. 
"Winwood  Reade  informs  me  that  the  negroes  of  West  Africa  often 
commit  smcide.  It  is  well  known  how  common  it  was  amongst  the 
miserable  aborigines  of  South  America,  after  the  Spanish   conquest. 


114  THE  DESCENT  OF  MAN. 

the  courage  displayed,  as  an  honorable  act;  and  it  is  still  prac- 
ticed by  some  semi-civilized  and  savage  nations  without  reproach, 
for  it  does  not  obviously  concern  others  of  the  tribe.  It  has  been 
recorded  that  an  Indian  Thug  conscientiously  regretted  that  he 
had  not  robbed  and  strangled  as  many  travelers  as  did  his  father 
before  him.  In  a  rude  state  of  civilization  the  robbery  of  strang- 
ers is,  indeed,  generally  considered  as  honorable. 

Slavery,  although  in  some  ways  beneficial  during  ancient 
times,"*  is  a  great  crime;  yet  it  was  not  so  regarded  until  quite 
recently,  even  by  the  most  civilized  nations.  And  this  was  espe- 
cially the  case,  because  the  slaves  belonged  in  general  to  a  race 
different  from  that  of  their  masters.  As  barbarians  do  not  regard 
the  opinion  of  their  women,  wives  are  commonly  treated  like 
slaves.  Most  savages  are  utterly  indifferent  to  the  sufferings  of 
strangers,  or  even  delight  in  witnessing  them.  It  is  well  known 
that  the  women  and  children  of  the  North-American  Indians 
aided  in  torturing  their  enemies.  Some  savages  take  a  horrid 
pleasure  in  cruelty  to  animals,^'^  and  humanity  is  an  unknown 
virtue.  Nevertheless,  besides  the  family  affections,  kindness  is 
common,  especially  during  sickness,  between  the  members  of 
the  same  tribe,  and  is  sometimes  extended  beyond  these  limits. 
Mungo  Park's  touching  account  of  the  kindness  of  the  negro 
women  of  the  interior  to  him  is  well  known.  Many  instances 
could  be  given  of  the  noble  fidelity  of  savages  towards  each  other, 
but  not  to  strangers;  common  experience  justifies  the  maxim 
of  the  Spaniard,  "Never,  never  trust  an  Indian."  There  cannot 
be  fidelity  without  truth;  and  this  fundamental  virtue  is  not 
rare  between  the  members  of  the  same  tribe:  thus  Mungo  Park 
heard  the  negro  women  teaching  their  young  children  to  love 
the  truth.  This,  again,  is  one  of  the  virtues  which  becomes  so 
deeply  rooted  in  the  mind,  that  it  is  sometimes  practiced  by  sav- 
ages, even  at  a  high  cost,  towards  strangers;  but  to  lie  to  your 
enemy  has  rarely  been  thought  a  sin,  as  the  history  of  modern 
diplomacy  too  plainly  shows.  As  soon  as  a  tribe  has  a  recognized 
leader,  disobedience  becomes  a  crime,  and  even  abject  submission 
is  looked  at  as  a  sacred  virtue. 

As  during  rude  times  no  man  can  be  useful  or  faithful  to  his 
tribe  without  courage,  this  quality  has  universally  been  placed 
in  the  highest  rank;  and  although  in  civilized  countries  a  good 
yet  timid  man  may  be  far  more  useful  to  the  community  than  a 
brave  one,  we  cannot  help  instinctively  honoring  the  latter  above 

For  New  Zealand,  see  the  voyage  of  the  "Novara,"  and  for  the  Aleu- 
tian Islands,  Muller,  as  quoted  by  Houzeau,  'Les  Facultes  Mentales,' 
&c.,  torn.  ii.  p.  136. 

34  See  Mr.  Bag-ehot,  'Physics  and  Politics,'  1872,  p.  72. 

35  See,  for  instance,  Mr.  Hamilton's  account  of  the  Kaffirs,  'Anthro- 
pological Review,'  1870,  p.  xv. 


MORAL   SENSE.  115 

a  coward,  however  benevolent.  Prudence,  on  the  other  hand, 
which  does  not  concern  the  welfare  of  others,  though  a  very  use- 
ful virtue,  has  never  been  highly  esteemed.  As  no  man  can 
practice  the  virtues  necessary  for  the  welfare  of  his  tribe  without 
self-sacrifice,  self-command,  and  the  power  of  endurance,  these 
qualities  have  been  at  all  times  highly  and  most  justly  valued. 
The  American  savage  voluntarily  submits  to  the  most  horrid 
tortures  without  a  groan,  to  prove  and  strengthen  his  fortitude 
and  courage;  and  we  cannot  help  admiring  him,  or  even  an  In- 
dian Fakir,  who,  from  a  foolish  religious  motive,  swings  sus- 
pended by  a  hook  buried  in  his  flesh. 

The  other  so  called  self-regarding  virtues,  which  do  not  ob- 
viously, though  they  may  really,  afiect  the  welfare  of  the  tribe, 
have  never  been  esteemed  by  savages,  though  now  highly  appre- 
ciated by  civilized  nations.  The  greatest  intemperance  is  no 
reproach  with  savages.  Utter  licentiousness,  and  unnatural 
crimes,  prevail  to  an  astounding  extent.^^  As  soon,  however,  as 
marriage,  whether  polygamous,  or  monogamous,  becomes  com- 
mon, jealousy  will  lead  to  the  inculcation  of  female  virtue;  and 
this,  being  honored,  will  tend  to  spread  to  the  unmarried  females. 
How  slowly  it  spreads  to  the  male  sex,  we  see  at  the  present  da3\ 
Chastity  eminently  requires  self-command;  therefore  it  has  been 
honored  from  a  very  early  period  in  the  moral  history  of  civilized 
man.  As  a  consequence  of  this,  the  senseless  practice  of  celibacy 
has  been  ranked  from  a  remote  period  as  a  virtue.^'  The  hatred 
of  indecency,  which  appears  to  us  so  natural  as  to  be  thought  in- 
nate, and  which  is  so  valuable  an  aid  to  chastity,  is  a  modern 
virtue,  appertaining  exclusively,  as  Sir  G.  Staunton  remarks,'^  to 
civilized  life.  This  is  shown  by  the  ancient  religious  rites  of 
various  nations,  by  the  drawings  on  the  w^alls  of  Pompeii,  and 
by  the  practices  of  many  savages. 

We  have  now  seen  that  actions  are  regarded  by  savages,  and 
were  probably  so  regarded  by  primaeval  man,  as  good  or  bad, 
solely  as  they  obviously  affect  the  welfare  of  the  tribe,— not  that 
of  the  species,  nor  that  of  an  individual  member  of  the  tribe. 
This  conclusion  agrees  well  with  the  belief  that  the  so-called 
moral  sense  is  aboriginally  derived  from  the  social  instincts,  for 
both  relate  at  first  exclusively  to  the  community.  The  chief 
causes  of  the  low  morality  of  savages,  as  judged  by  our  standard, 
are,  firstly,  the  confinement  of  sympathy  to  the  same  tribe.  Sec- 
ondly, powers  of  reasoning  insufficient  to  recognize  the  bearing 
of  many  virtues,  especially  of  the  self-regarding  virtues,  on  the 


38  Mr.  M'Lennan  has  given  ('Primitive  Marriage,'  1865,  p.  176)  a  good 
collection  of  facts  on  this  head. 
3'^  Lecky,   'History  of  European  Morals,'  vol.  i.  1869,  p.  109. 
^  'Embassy  to  China,'  vol.  ii.  p.  348. 


116  THE   DESCENT  OF   MAN. 

general  welfare  of  the  tribe.  Savages,  for  instance,  fail  to  trace 
the  multiplied  evils  consequent  on  a  want  of  temperance,  chas- 
tity, &c.  And,  thirdly,  weak  power  of  self  command;  for  this 
power  has  not  heen  strengthened  through  long-continued,  per- 
haps inherited,  habit,  instruction  and  religion. 

I  have  entered  into  the  above  details  on  the  immorality  of 
savages,^^  because  some  authors  have  recently  taken  a  high  view 
of  their  moral  nature,  or  have  attributed  most  of  their  crimes  to 
mistaken  benevolence. *°  These  authors  appear  to  rest  their  con- 
clusion on  savages  possessing  those  virtues  which  are  serviceable, 
or  even  necessary,  for  the  existence  of  the  family  and  of  the 
tribe, — qualities  which  they  undoubtedly  do  possess,  and  often  in 
a  high  degree. 


Concluding  Bemarks. — It  was  assumed  formerly  by  philoso- 
phers of  the  derivative*^  school  of  morals  that  the  foundation  of 
morality  lay  in  a  form  of  Selfishness;  but  more  recently  the 
"Greatest  happiness  principle"  has  been  brought  prominently 
forward.  It  is,  however,  more  correct  to  speak  of  the  latter  prin- 
ciple as  the  standard,  and  not  as  the  motive  of  conduct.  Never- 
theless, all  the  authors  whose  works  I  have  consulted,  with  a  few 
exceptions,^  write  as  if  there  must  be  a  distinct  motive  for  every 
action,  and  that  this  must  be  associated  with  some  pleasure  or 
displeasure.  But  man  seems  often  to  act  impulsively,  that  is 
from  instinct  or  long  habit,  without  any  consciousness  of  pleasure, 

3»  See  on  this  subject  copious  evidence  in  Chap.  vii.  of  Sir  J.  Lub- 
bock, 'Oi-igin  of  Civilization,'  1870. 

^°  For  instance,  Lecky,  'Hist.  European  Morals,'  vol.  i.  p.  124. 

^  This  term  is  used  in  an  able  article  in  the  'Westminster  Review,' 
Oct.  1869,  p.  498.  For  the  "Greatest  happiness  principle,"  see  J.  S.  Mill, 
'Utilitarianism,'  p.  17. 

^  Mill  recognizes  ('System  of  Logic,*  vol.  ii.  p.  422)  in  the  clearest 
manner,  that  actions  may  be  performed  through  habit  without  the 
anticipation  of  pleasure.  Mr.  H.  Sidgwick  also  in  his  Essay  on  Pleas- 
ure and  Desire  ('The  Contemporary  Review,' April  1872,  p.  671),  remarks: 
"To  sum  up,  in  contravention  of  the  doctrine  that  our  conscious  ac- 
"tive  impulses  are  always  directed  towards  the  production  of  agreeable 
"sensations  in  ourselves,  I  would  maintain  that  we  find  everywhere  in 
"consciousness  extra-regarding  impulse,  directed  towards  something 
"that  is  not  pleasure;  that  in  many  cases  the  impulse  is  so  far  incom- 
"patible  with  the  self -regarding  that  the  two  do  not  easily  co-exist  in 
"the  same  moment  of  consciousness."  A  dim  feeling  that  our  impulses 
do  not  by  any  means  always  arise  from  any  contemporaneous  or  an- 
ticipated pleasure,  has,  I  cannot  but  think,  been  one  chief  cause  of  the 
acceptance  of  the  intuitive  theory  of  morality,  and  of  the  rejection  of 
the  utilitarian  or  "Greatest  happiness"  theory.  With  respect  to  the 
latter  theory,  the  standard  and  the  motive  of  conduct  have  no  doubt 
often  been  confused,  but  they  are  really  in  some  degree   blended. 


MORAL   SENSE.  11? 

in  the  same  manner  as  does  probably  a  bee  or  ant,  when  it 
blindly  follows  its  instincts.  Under  circumstances  of  extreme 
peril,  as  during  a  fire,  when  a  man  endeavors  to  save  a  fellow- 
creature  without  a  moment's  hesitation,  he  can  hardly  feel  pleas- 
ure; and  still  less  has  he  time  to  reflect  on  the  dissatisfaction 
which  he  might  subsequently  experience  if  he  did  not  make  the 
attempt.  Should  he  afterwards  reflect  over  his  own  conduct,  he 
would  feel  that  there  lies  within  him  an  impulsive  power  widely 
different  from  a  search  after  pleasure  or  happiness;  and  this 
seems  to  be  the  deeply  planted  social  instinct. 

In  the  case  of  the  lower  animals  it  seems  much  more  appro- 
priate to  speak  of  their  social  instincts,  as  having  been  developed 
for  the  general  good  rather  than  for  the  general  happiness  of  the 
species.  The  term,  general  good,  may  be  deflned  as  the  rearing 
of  the  greatest  number  of  individuals  in  full  vigor  and  health, 
with  all  their  faculties  perfect,  under  the  conditions  to  which 
they  are  subjected.  As  the  social  instincts  both  of  man  and  the 
lower  animals  have  no  doubt  been  developed  by  nearly  the  same 
steps,  it  would  be  advisable,  if  found  practicable,  to  use  the  same 
definition  in  both  cases,  and  to  take  as  the  standard  of  morality, 
the  general  good  or  welfare  of  the  community,  rather  than  the 
general  happiness;  but  this  definition  would  perhaps  require  some 
limitation  on  account  of  political  ethics. 

When  a  man  risks  his  life  to  save  that  of  a  fellow-creature,  it 
seems  also  more  correct  to  say  that  he  acts  for  the  general  good, 
rather  than  for  the  general  happiness  of  mankind.  No  doubt 
the  welfare  and  the  happiness  of  the  individual  usually  coincide; 
and  a  contented,  happy  tribe  will  flourish  better  than  one  that 
is  discontented  and  unhappy.  We  have  seen  that  even  at  an 
early  period  in  the  history  of  man,  the  expressed  wishes  of  the 
community  will  have  naturally  influenced  to  a  large  extent  the 
conduct  of  each  member;  and  as  all  wish  for  happiness,  the 
"greatest  happiness  principle"  will  have  become  a  most  important 
secondary  guide  and  object;  the  social  instinct,  however,  together 
with  sympathy  (which  leads  to  our  regarding  the  approbation 
and  disapprobation  of  others),  having  served  as  the  primary  im- 
pulse and  guide.  Thus  the  reproach  is  removed  of  laying  the 
foundation  of  the  noblest  part  of  our  nature  in  the  base  principle 
of  selflshness;  unless,  indeed,  the  satisfaction  which  every  animal 
feels,  when  it  follows  its  proper  instincts,  and  the  dissatisfaction 
felt  when  prevented,  be  called  selfish. 

The  wishes  and  opinions  of  the  members  of  the  same  com- 
munity, expressed  at  first  orally,  but  later  by  writing  also,  either 
form  the  sole  guides  of  our  conduct,  or  greatly  reinforce  the 
social  instincts;  such  opinions,  however,  have  sometimes  a  ten- 
dency directly  opposed  to  these  instincts.  This  latter  fact  is 
well  exemplified  by  the  Law  of  Honor,  that  is,  the  law  of  the 

O 


118  THE  DESCENT  OF  MAN. 

opinion  of  our  equals,  and  not  of  all  our  countrymen.  The  breach 
of  this  law,  even  when  the  breach  is  known  to  be  strictly  ac- 
cordant with  true  morality,  has  caused  many  a  man  more  agony 
than  a  real  crime.  We  recognize  the  same  influence  in  the  burn- 
ing sense  of  shame  which  most  of  us  have  felt,  even  after  the 
interval  of  years,  when  calling  to  mind  some  accidental  breach 
of  a  trifling,  though  fixed,  rule  of  etiquette.  The  judgment  of 
the  community  will  generally  be  guided  by  some  rude  experience 
of  what  is  best  in  the  long  run  for  all  the  members;  but  this 
judgment  will  not  rarely  err  from  ignorance  and  weak  powers  of 
reasoning.  Hence  the  strangest  customs  and  superstitions,  in 
complete  opposition  to  the  true  welfare  and  happiness  of  man- 
kind, have  become  all-powerful  throughout  the  world.  We  see 
this  in  the  horror  felt  by  a  Hindoo  who  breaks  his  caste,  and 
in  many  other  such  cases.  It  would  be  difficult  to  distinguish 
between  the  remorse  felt  by  a  Hindoo  who  has  yielded  to  the 
temptation  of  eating  unclean  food,  from  that  felt  after  commit- 
ting a  theft;    but  the  former  would  probably  be  the  more  severe. 

How  so  many  absurd  rules  of  conduct,  as  well  as  so  many  absurd 
religious  beliefs,  have  originated,  we  do  not  know;  nor  how 
it  is  that  they  have  become,  in  all  quarters  of  the  world,  so 
deeply  impressed  on  the  mind  of  men;  but  it  is  worthy  of  re- 
mark that  a  belief  constantly  inculcated  during  the  early  years  of 
life,  whilst  the  brain  is  impressible,  appears  to  acquire  almost 
the  nature  of  an  instinct;  and  the  very  essence  of  an  instinct  is 
that  it  is  followed  independently  of  reason.  Neither  can  we  say 
why  certain  admirable  virtues,  such  as  the  love  of  truth,  are 
much  more  highly  appreciated  by  some  savage  tribes  than  by 
others;*^  nor,  again,  why  similar  differences  prevail  even  amongst 
highly  civilized  nations.  Knowing  how  firmly  fixed  many  strange 
customs  and  superstitions  have  become,  we  need  feel  no  surprise 
that  the  self-regarding  virtues,  supported  as  they  are  by  reason, 
should  now  appear  to  us  so  natural  as  to  be  thought  innate,  al- 
though they  were  not  valued  by  man  in  his  early  condition. 

Notwithstanding  many  sources  of  doubt,  man  can  generally 
and  readily  distinguish  between  the  higher  and  lower  moral 
rules.  The  higher  are  founded  on  the  social  instincts,  and  relate 
to  the  welfare  of  others.  They  are  supported  by  the  approbation 
of  our  fellow-men  and  by  reason.  The  lower  rules,  though  some 
of  them,  when  implying  self-sacrifice  hardly  deserve  to  be  called 
lower,  relate  chiefly  to  self,  and  arise  from  public  opinion,  ma- 
tured by  experience  and  cultivation;  for  they  are  not  practiced 
by  rude  tribes. 

*3  Good  instances  are  g-iven  by  Mr.  Wallace  in  'Scientific  Opinion,* 
Sept.  15,  1869;  and  more  fully  in  his  'Contributions  to  the  Theory  of 
Natural  Selection,'  1870,  p.  353. 


MORAL    SENSE.  119 

As  man  advances  in  civilization,  and  small  tribes  are  united 
into  larger  communities,  the  simplest  reason  would  tell  each 
individual  that  he  ought  to  extend  his  social  instincts  and  sym- 
pathies to  all  the  members  of  the  same  nation,  though  personally 
unknown  to  him.  This  point  being  once  reached,  there  is  only  an 
artificial  barrier  to  prevent  his  sympathies  extending  to  the  men 
of  all  nations  and  races.  If,  indeed,  such  men  are  separated  from 
tim  by  great  differences  in  appearance  or  habits,  experience  un- 
fortunately shows  us  how  long  it  is,  before  we  look  at  them  as 
our  fellow-creatures.  Sympathy  beyond  the  confines  of  man,  that 
is,  humanity  to  the  lower  animals,  seems  to  be  one  of  the  latest 
moral  acquisitions.  It  is  apparently  unfelt  by  savages,  except 
towards  their  pets.  How  little  the  old  Romans  knew  of  it  is 
shown  by  their  abhorrent  gladiatorial  exhibitions.  The  very  idea 
of  humanity,  as  far  as  I  could  observe,  was  new  to  most  of  the 
Gauchos  of  the  Pampas.  This  virtue,  one  of  the  noblest  with 
which  man  is  endowed,  seems  to  arise  incidentally  from  our  sym- 
pathies becoming  more  tender  and  more  widely  diffused,  until 
they  are  extended  to  all  sentient  beings.  As  soon  as  this  virtue 
is  honored  and  practiced  by  some  few  men,  it  spreads  through 
instruction  and  example  to  the  young,  and  eventually  becomes 
incorporated  in  public  opinion. 

The  highest  possible  stage  in  moral  culture  is  when  we  recog- 
nize that  we  ought  to  control  our  thoughts,  and  "not  even  in 
"inmost  thought  to  think  again  the  sins  that  made  the  past  so 
"pleasant  to  us."^*  Whatever  makes  any  bad  action  familiar  to 
the  mind,  renders  its  performance  by  so  much  the  easier.  As 
Marcus  Aurelius  long  ago  said,  "Such  as  are  thy  habitual 
"thoughts,  such  also  will  be  the  character  of  thy  mind;  for  the 
"soul  is  dyed  by  the  thoughts."*^ 

Our  great  philosopher,  Herbert  Spencer,  has  recently  explained 
his  views  on  the  moral  sense.  He  says,**'  "I  believe  that  the 
"experiences  of  utility  organized  and  consolidated  through  all 
"past  generations  of  the  human  race,  h^ave  been  producing  cor- 
"responding  modifications,  which,  by  continued  transmission  and 
"accumulation,  have  become  in  us  certain  faculties  of  moral 
"intuition — certain  emotions  responding  to  right  and  wrong  con- 
"ducfc,  which  have  no  apparent  basis  in  the  individual  experiences 
"of  utility."  There  is  not  the  least  inherent  improbability,  as  it 
seems  to  me,  in  virtuous  tendencies  being  more  or  less  strongly 
inherited;  for,  not  to  mention  the  various  dispositions  and  habits 
transmitted  by  many  of  our  domestic  animals  to  their  offspring, 

**  Tennyson,  'Idylls  of  the  King,'  p.  244. 

45  'The  Thoughts  of  the  Emperor  M.  Aurelius  Antoninus,'  Eng.  trans- 
lat.,  2nd  edit.,  1869,  p.  112.    Marcus  Aurelius  was  born  A.  D.  121. 
*«  Letter  to  Mr.  Mill  in  Bain's  'Mental  and  Moral  Science,'  18o8,  p.  722. 


120  THE  DESCENT  OP  MAN. 

I  have  heard  of  authentic  cases  in  which  a  desire  to  steal  and 
a  tendency  to  lie  appeared  to  run  in  families  of  the  upper  ranks; 
and  as  stealing  is  a  rare  crime  in  the  wealthy  classes,  we  can 
hardly  account  by  accidental  coincidence  for  the  tendency  occur- 
ring in  two  or  three  members  of  the  same  family.  If  bad  ten- 
dencies are  transmitted,  it  is  probable  that  good  ones  are  like- 
wise transmitted.  That  the  state  of  the  body  by  affecting  the 
brain,  has  great  influence  on  the  moral  tendencies  is  known  to 
most  of  those  who  have  suffered  from  chronic  derangements  of 
the  digestion  or  liver.  The  same  fact  is  likewise  shown  by  the 
"perversion  or  destruction  of  the  moral  sense  being  often  one  of 
^'the  earliest  symptoms  of  mental  derangement;"*^  and  insanity 
is  notoriously  often  inherited.  Except  through  the  principle  of 
the  transmission  of  moral  tendencies,  we  cannot  understand  the 
differences  believed  to  exist  in  this  respect  between  the  various 
races  of  mankind. 

Even  the  partial  transmission  of  virtuous  tendencies  would 
be  an  immense  assistance  to  the  primary  impulse  derived  directly 
and  indirectly  from  the  social  instincts.  Admitting  for  a  moment 
that  virtuous  tendencies  are  inherited,  it  appears  probable,  at 
least  in  such  cases  as  chastity,  temperance,  humanity  to  animals, 
&c.,  that  they  become  first  impressed  on  the  mental  organization 
through  habit,  instruction  and  example,  continuied  during  sev- 
eral generations  in  the  same  family,  and  in  a  quite  subordinate 
degree,  or  not  at  all,  by  the  individuals  possessing  such  virtues 
having  succeeded  best  in  the  struggle  for  life.  My  chief  source 
of  doubt  with  respect  to  any  such  inheritance,  is  that  senseless 
customs,  superstitions,  and  tastes,  such  as  the  horror  of  a  Hindoo 
for  unclean  food,  ought  on  the  same  principle  to  be  transmitted. 
I  have  not  met  with  any  evidence  in  support  of  the  transmission 
of  superstitious  customs  or  senseless  habits,  although  in  itself  it 
is  perhaps  not  less  probable  than  that  animals  should  acquire 
inherited  tastes  for  certain  kinds  of  food  or  fear  of  certain  foes. 

Finally  the  social  instincts,  which  no  doubt  were  acquired  by 
man  as  by  the  lower  animals  for  the  good  of  the  community, 
will  from  the  first  have  given  to  him  some  wish  to  aid  his 
fellows,  some  feeling  of  sympathy,  and  have  compelled  him  to 
regard  their  approbation  and  disapprobation.  Such  impulses 
!  will  have  served  him  at  a  very  early  period  as  a  rude  rule  of 
\  right  and  wrong.  But  as  man  gradually  advanced  in  intellectual 
power,  and  was  enabled  to  trace  the  more  remote  consequences 
of  his  actions;  as  he  acquired  sufiicient  knowledge  to  reject 
baneful  customs  and  superstitions;  as  he  regarded  more  and 
more,  not  only  the  welfare,  but  the  happiness  of  his  fellow-men; 


*7  Maudsley,  'Body  and  Mind,'  1870,  p.  60. 


SUMMARY.  121     -^ 

as  from  habit,  following  on  beneficial  experience,  instruction  1 
and  example,  his  sympathies  became  more  tender  and  widely  | 
diffused,  extending  to  men  of  all  races,  to  the  imbecile,  maimed,  | 
and  other  useless  members  of  society,  and  finally  to  the  lower  \ 
animals, — so  would  the  standard  of  his  morality  rise  higher  and  ] 
higher.  And  it  is  admitted  by  moralists  of  the  derivative  school 
and  by  some  intuitionists,  that  the  standard  of  morality  has  ; 
risen  since  an  early  period  in  the  history  of  man.*^ 

As  a  struggle  may  sometimes  be  seen  going  on  between  the 
various  instincts  of  the  lower  animals,  it  is  not  surprising  that 
there  should  be  a  struggle  in  man  between  his  social  instincts, 
with  their  derived  virtues,  and  his  lower,  though  momentarily 
stronger  impulses  or  desires.  This,  as  Mr.  Galton  *^  has  remarked, 
is  all  the  less  surprising,  as  man  has  emerged  from  a  state  of 
barbarism  within  a  comparatively  recent  period.  After  having 
yielded  to  some  temptation  we  feel  a  sense  of  dissatisfaction, 
shame,  repentance,  or  remorse,  analogous  to  the  feelings  caused 
by  other  powerful  instincts  or  desires,  when  left  unsatisfied  or 
baulked.  We  compare  the  weakened  impression  of  a  past  tempta- 
tion with  the  ever  present  social  instincts,  or  with  habits,  gained 
in  early  youth  and  strengthened  during  our  whole  lives,  until 
they  have  become  almost  as  strong  as  instincts.  If  with  the 
temptation  still  before  us  we  do  not  yield,  it  is  because  either 
the  social  instinct  or  some  custom  is  at  the  moment  predominant, 
or  because  we  have  learnt  that  it  will  appear  to  us  hereafter  the 
stronger,  when  compared  with  the  weakened  impression  of  the  , 
temptation,  and  we  realize  that  its  violation  would  cause  us  suf-  i 
fering.  Looking  to  future  generations,  there  is  no  cause  to  fear  J 
that  the  social  instincts  will  grow  weaker,  and  we  may  expect  that  "I 


virtuous  habits  will  grow  stronger,  becoming  perhaps  fixed  by  | 
inheritance.  In  this  case  the  struggle  between  our  higher  and  f 
lower  impulses  will  be  less  severe,  and  virtue  will  be  triumphant. 


Summary  of  the  last  two  Clia'pters. — There  can  be  no  doubt  that 
the  difference  between  the  mind  of  the  lowest  man  and  that  of 
the  highest  animal  is  immense.  An  anthropomorphous  ape,  if 
he  could  take  a  dispassionate  view  of  his  own  case,  would  admit 
that  though  he  could  form  an  artful  plan  to  plunder  a  garden — 
though  he  could  use  stones   for  fighting  or  for  breaking  open 

*8A  writer  in  the  'North  British  Review'  (July,  1869,  p.  531),  well 
capable  of  forming  a  sound  judgment,  expresses  himself  strongly  in 
favor  of  this  conclusion.  Mr.  Lecky  ('Hist,  of  Morals,'  vol.  i.  p.  143) 
seems  to  a  certain  extent  to  coincide  therein. 

*®  See  his  remarkable  work  on  'Hereditary  Genius,'  1869,  p.  349.  The 
Duke  of  Argyll  ('Primeval  Man,'  1869,  p.  188)  has  some  good  remarks  on 
the  contest  in  man's  nature  between  right  and  wrong. 


122  THE  DESCENT  OF  MAK. 

nuts,  yet  that  the  thought  of  fashioning  a  stone  into  a  tool  was 
quite  beyond  his  scope.  Still  less,  as  he  would  admit,  could  he 
follow  out  a  train  of  metaphysical  reasoning,  or  solve  a  mathe- 
matical problem,  or  reflect  on  God,  or  admire  a  grand  natural 
scene.  Some  apes,  however,  would  probably  declare  that  they 
could  and  did  admire  the  beauty  of  the  colored  skin  and  fur  of 
their  partners  in  marriage.  They  would  admit,  that  though  they 
could  make  other  apes  understand  by  cries  some  of  their  per- 
ceptions and  simpler  wants,  the  notion  of  expressing  definite  ideas 
by  definite  sounds  had  never  crossed  their  minds.  They  might 
insist  that  they  were  ready  to  aid  their  fellow-apes  of  the  same 
troop  in  many  ways,  to  risk  their  lives  for  them,  and  to  take 
charge  of  their  orphans;  but  they  would  be  forced  to  acknowl- 
edge that  disinterested  love  for  all  living  creatures,  the  most 
noble  attribute  of  man,  was  quite  beyond  their  comprehension. 

Nevertheless  the  difference  in  mind  between  man  and  the 
higher  animals,  great  as  it  is,  certainly  is  one  of  degree  and  not 
of  kind.  We  have  seen  that  the  senses  and  intuitions,  the 
various  emotions  and  faculties,  such  as  love,  memory,  attention, 
curiosity,  imitation,  reason,  &c.,  of  which  man  boasts,  may  be 
found  in  an  incipient,  or  even  sometimes  in  a  well-developed 
condition,  in  the  lower  animals.  They  are  also  capable  of  some 
inherited  improvement,  as  we  see  in  the  domestic  dog  compared 
with  the  wolf  or  jackal.  If  it  could  be  proved  that  certain  high 
mental  powers,  such  as  the  formation  of  general  concepts,  self- 
consciousness,  &c.,  were  absolutely  peculiar  to  man,  which  seems 
extremely  doubtful,  it  is  not  improbable  that  these  qualities  are 
merely  the  incidental  results  of  other  highly-advanced  intellec- 
tual faculties;  and  these  again  mainly  the  result  of  the  continued 
use  of  a  perfect  language.  At  what  age  does  the  new-born  in- 
fant possess  the  power  of  abstraction,  or  become  self-conscious, 
land  reflect  on  its  own  existence?  We  cannot  answer;  nor  can 
I  we  answer  in  regard  to  the  ascending  organic  scale.  The  half- 
';•  art,  half-instinct  of  language  still  bears  the  stamp  of  its  gradual 
evolution.  The  ennobling  belief  in  God  is  not  universal  with 
man;  and  the  belief  in  spiritual  agencies  naturally  follows  from 
other  mental  powers.  The  moral  sense  perhaps  affords  the  best 
and  highest  distinction  between  man  and  the  lower  animals;  but 
I  need  say  nothing  on  this  head,  as  I  have  so  lately  endeavored 
to  show  that  the  social  instincts, — the  prime  principle  of  man's 
moral  constitution^^ — with  the  aid  of  active  intellectual  powers 
and  the  effects  of  habit,  naturally  lead  to  the  golden  rule,  "As  ye 
"would  that  men  should  do  to  you,  do  ye  to  them  likewise;"  and 
this  lies  at  the  foundation  of  morality. 

50  'The  Thoughts  of  Marcus  Aurelius,'  &c.,  p.  138. 


SUMMARY.  123 

In  the  next  chapter  I  shall  make  some  few  remarks  on  the 
probable  steps  and  means  by  which  the  several  mental  and  moral 
faculties  of  man  have  been  gradually  evolved.  That  such  evolu- 
tion is  at  least  possible,  ought  not  to  be  denied,  for  we  daily  see 
these  faculties  developing  in  every  infant;  and  we  may  trace  a 
perfect  gradation  from  the  mind  of  an  utter  idiot,  lower  than 
that  of  an  animal  low  in  the  scale,  to  the  mind  of  a  Newton. 


12i  THE  DESCENT  OF  MAN. 


CHAPTER  V. 

ON  THE  DEVELOPMENT  OF  THE  INTELLECTUAL  AND  MORAL 
FACULTIES  DURING  PRIMEVAL  AND  CIVILIZED   TIMES. 

Advancement  of  the  intellectual  powers  through  natural  selection — 
Importance  of  imitation— Social  and  moral  faculties— Their  devel- 
opment within  the  limits  of  the  same  tribe — Natural  selection  as 
affecting  civilized  nations— Evidence  that  civilized  nations  were 
once  barbarous. 

The  subjects  to  be  discussed  in  this  chapter  are  of  the  highest 
interest,  but  are  treated  by  me  in  an  imperfect  and  fragmentary 
manner.  Mr.  Wallace,  in  an  admirable  paper  before  referred  to,* 
argues  that  man,  after  he  had  partially  acquired  those  intellec- 
tual and  moral  faculties  which  distinguish  him  from  the  lower 
animals,  would  have  been  but  little  liable  to  bodily  modifica- 
tions through  natural  selection  or  any  other  means.  For  man  is 
enabled  through  his  mental  faculties  "to  keep  with  an  unchanged 
"body  in  harmony  with  the  changing  universe."  He  has  great 
power  of  adapting  his  habits  to  new  conditions  of  life.  He  in- 
vents weapons,  tools,  and  various  stratagems  to  procure  food  and 
to  defend  himself.  When  he  migrates  into  a  colder  climate  he 
uses  clothes,  builds  sheds,  and  makes  fires;  and  by  the  aid  of 
fire  cooks  food  otherwise  indigestible.  He  aids  his  fellow-men  in 
many  ways,  and  anticipates  future  events.  Even  at  a  remote 
period  he  practiced  some  division  of  labor. 

The  lower  animals,  on  the  other  hand,  must  have  their  bodily 
structure  modified  in  order  to  survive  under  greatly  changed 
conditions.  They  must  be  rendered  stronger,  or  acquire  more 
effective  teeth  or  claws,  for  defense  against  new  enemies;  or 
they  must  be  reduced  in  size,  so  as  to  escape  detection  and  dan- 
ger. When  they  migrate  into  a  colder  climate,  they  must  become 
clothed  with  thicker  fur,  or  have  their  constitutions  altered.  If 
they  fail  to  be  thus  modified,  they  will  cease  to  exist. 

The  case,  however,  is  widely  different,  as  Mr.  Wallace  has  with 
justice  insisted,  in  relation  to  the  intellectual  and  moral  faculties 


1  'Anthropological  Review,'  May,  1864,  p.  clviii. 


INTELLECTUAL   FACULTIES.  125 

of  man.  These  faculties  are  variable;  and  we  have  every  reason 
to  believe  that  the  variations  tend  to  be  inherited.  Therefore,  if 
they  were  formerly  of  high  importance  to  primeval  man  and  to 
his  ape-like  progenitors,  they  would  have  been  perfected  or  ad- 
vanced through  natural  selection.  Of  the  high  importance  of  the 
intellectual  faculties  there  can  be  no  doubt,  for  man  mainly  owes 
to  them  his  predominant  position  in  the  world.  We  can  see, 
that  in  the  rudest  state  of  society,  the  individuals  who  were  the 
most  sagacious,  who  invented  and  used  the  best  weapons  or 
traps,  and  who  were  best  able  to  defend  themselves,  would  rear 
the  greatest  number  of  offspring.  The  tribes,  which  included 
the  largest  number  of  men  thus  endowed,  would  increase  in 
number  and  supplant  other  tribes.  Numbers  depend  primarily 
on  the  means  of  subsistence,  and  this  depends  partly  on  the  phys- 
ical nature  of  the  country,  but  in  a  much  higher  degree  on  the 
arts  which  are  there  practiced.  As  a  tribe  increases  and  is  vic- 
torious, it  is  often  still  further  increased  by  the  absorption  of 
other  tribes.^  The  stature  and  strength  of  the  men  of  a  tribe 
are  likewise  of  some  importance  for  its  success,  and  these  depend 
in  part  on  the  nature  and  amount  of  the  food  which  can  be 
obtained.  In  Europe  the  men  of  the  Bronze  period  were  sup- 
planted by  a  race  more  powerful,  and,  judging  from  their  sword- 
handles,  with  larger  han<ls;^  but  their  success  was  probably  still 
more  due  to  their  superiority  in  the  arts. 

All  that  we  know  about  savages,  or  may  infer  from  their  tra- 
ditions and  from  old  monuments,  the  history  of  which  is  quite 
forgotten  by  the  present  inhabitants,  show  that  from  the  remotest 
times  successful  tribes  have  supplanted  other  tribes.  Relics  of 
extinct  or  forgotten  tribes  have  been  discovered  throughout  the 
civilized  regions  of  the  earth,  on  the  wild  plains  of  America,  and 
on  the  isolated  islands  in  the  Pacific  Ocean.  At  the  present  day 
civilized  nations  are  everywhere  supplanting  barbarous  nations, 
excepting  where  the  climate  opposes  a  deadly  barrier;  and  they 
succeed  mainly,  though  not  exclusively,  through  their  arts,  which 
are  the  products  of  the  intellect.  |^It  is,  therefore,  highly  probable 
that  with  mankind  the  intellecthal  faculties  have  been  mainly 
and  gradually  perfected  through  natural  selection;  and  this  con- 
clusion is  sufficient  for  our  purpose.  Undoubtedly  it  would  be 
interesting  to  trace  the  development  of  each  separate  faculty 
from  the  state  in  which  it  exists  in  the  lower  animals  to  that  in 
which  it  exists  in  man;  but  neither  my  ability  nor  knowledge 
permits  the  attempt. 

2  After  a  time  the  members  or  tribes  which  are  absorbed  into  another 
tribe  assume,  as  Sir  Henry  Maine  remarks  ('Ancient  Law,'  1831,  p  7^), 
that  they  are  the  co-descendants  of  the  same  ancestors. 

»  Morlot,  'Soc.  Vaud.  Sc.  Nat.'  1860,  p.  294. 


126  THE  DESCENT  OF  MAN. 

It  deserves  notice  that,  as  soon  as  the  progenitors  of  man  be- 
came social  (and  this  probably  occurred  at  a  very  early  period), 
the  principle  of  imitation,  and  reason,  and  experience  would  have 
increased,  and  much  modified  the,  intellectual  powers  in  a  way, 
of  which  we  see  only  traces  in  the  lower  animals.  Apes  are 
much  given  to  imitation,  as  are  the  lowest  savages;  and  the 
simple  fact  previously  referred  to,  that  after  a  time  no  animal 
can  be  caught  in  the  same  place  by  the  same  sort  of  trap,  shows 
that  animals  learn  by  experience,  and  imitate  the  caution  of 
others.  Now,  if  some  one  man  in  a  tribe,  more  sagacious  than 
the  others,  invented  a  new  snare  or  weapon,  or  other  means  of 
attack  or  defense,  the  plainest  self-interest,  without  the  assistance 
of  much  reasoning  power,  would  prompt  the  other  members  to 
imitate  him;  and  all  would  thus  profit.  The  habitual  practice 
of  each  new  art  must  likewise  in  some  slight  degree  strengthen 
the  intellect.  If  the  new  invention  were  an  important  one,  the 
tribe  would  increase  in  number,  spread,  and  supplant  other  tribes. 
In  a  tribe  thus  rendered  more  numerous  there  would  always  be 
a  rather  greater  chance  of  the  birth  of  other  superior  and  in- 
ventive members.  If  such  men  left  children  to  inherit  their  men- 
tal superiority,  the  chance  of  the  birth  of  still  more  ingenious 
members  would  be  somewhat  better,  and  in  a  very  small  tribe 
decidedly  better.  Even  if  theyi  left  no  children,  the  tribe  would 
still  include  their  blood-relations;  and  it  has  been  ascertained 
by  agriculturists*  that  by  preserving  and  breeding  from  the 
family  of  an  animal,  which  when  slaughtered  was  found  to  be 
valuable,  the  desired  character  has  been  obtained. 

Turning  now  to  the  social  and  moral  faculties.  In  order  that 
primeval  men,  or  the  ape-like  progenitors  of  man,  should  become 
social,  they  must  have  acquired  the  same  instinctive  feelings, 
which  impel  other  animals  to  live  in  a  body;  and  they  no  doubt 
exhibited  the  same  general  disposition.  They  would  have  felt 
uneasy  when  separated  from  their  comrades,  for  whom  they 
would  have  felt  some  degree  of  love;  they  v/ould  have  v/arned 
each  other  of  danger,  and  have  given  mutual  aid  in  attack  or 
defense.  All  this  implies  some  degree  of  sympathy,  fidelity,  and 
courage.  Such  social  qualities,  the  paramount  importance  of 
which  to  the  lower  animals  is  disputed  by  no  one,  were  no  doubt 
acquired  by  the  progenitors  of  man  in  a  similar  manner,  namely, 
through  natural  selection,  aided  by  inherited  habit.  When  two 
'  tribes  of  primeval  man,  living  in  the  same  country,  came  into 
,  competition,  if  (other  circumstances  being  equal)  the  one  tribe 
f  included  a  great  number  of  courageous,  sympathetic  and  faithful 

*  I  have  given  instances  in  my  'Variation  of  Animals  under  Domesti- 
cation,' vol.  ii.  p.  196. 


MORAL    FACULTIES.  127 

members,  who  were  always  ready  to  warn  each  other  of  danger, 
to  aid  and  defend  each  other,  this  tribe  would  succeed  better  and 
conquer  the  other.  Let  it  be  borne  in  mind  how  all-important 
in  the  never-ceasing  wars  of  savages,  fidelity  and  courage  must 
be.  The  advantage  which  disciplined  soldiers  have  over  undis- 
ciplined hordes  follows  chiefly  from  the  confidence  which  each 
man  feels  in  his  comrades.  Obedience,  as  Mr.  Bagehot  has  well 
shown,^  is  of  the  highest  value,  for  any  form  of  government  is 
better  than  none.  Selfish  and  contentious  people  will  not  cohere, 
and  without  coherence  nothing  can  be  effected.  A  tribe  rich  in 
the  above  qualities  would  spread  and  be  victorious  over  other 
tribes:  but  in  the  course  of  time  it  would,  judging  from  all  past 
history,  be  in  its  turn  overcome  by  some  other  tribe  still  more 
highly  endowed.  Thus  the  social  and  moral  qualities  would  tend 
slowly  to  advance  and  be  diffused  throughout  the  world. 

But  it  may  be  asked,  how  within  the  limits  of  the  same  tribe 
did  a  large  number  of  members  first  become  endowed  with  these 
social  and  moral  qualities,  and  how  was  the  standard  of  ex- 
cellence raised?  It  is  extremely  doubtful  whether  the  offspring 
of  the  more  sympathetic  and  benevolent  parents,  or  of  those 
who  were  the  most  faithful  to  their  comrades,  would  be  reared 
in  greater  numbers  than  the  children  of  selfish  and  treacherous 
parents  belonging  to  the  same  tribe.  He  who  was  ready  to  sacri- 
fice his  life,  as  many  a  savage  has  been,  rather  than  betray  his 
comrades,  would  often  leave  no  offspring  to  inherit  his  noble 
nature.  The  bravest  men,  who  were  always  willing  to  come  to 
the  front  in  war,  and  who  freely  risked  their  lives  for  others, 
would  on  an  average  perish  in  larger  numbers  than  other  men. 
Therefore  it  hardly  seems  probable,  that  the  number  of  men 
gifted  with  such  virtues,  or  that  the  standard  of  their  excellence, 
could  be  increased  through  natural  selection,  that  is,  by  the 
survival  of  the  fittest;  for  we  are  not  here  speaking  of  one  tribe 
being  victorious  over  another. 

Although  the  circumstances,  leading  to  an  increase  in  the  num- 
ber of  those  thus  endowed  within  the  same  tribe,  are  too  complex 
to  be  clearly  followed  out,  we  can  trace  some  of  the  probable 
steps.  In  the  first  place,  as  the  reasoning  powers  and  foresight 
of  the  members  became  improved,  each  man  would  soon  learn 
that  if  he  aided  his  fellow-men,  he  would  commonly  receive  aid  in 
return.  From  this  low  motive  he  might  acquire  the  habit  of 
aiding  his  fellows;  and  the  habit  of  performing  benevolent  ac- 
tions certainly  strengthens  the  feeling  of  sympathy  which  gives 

*  See  a  remarkable  series  of  articles  on  'Physics  and  Politics'  in  the 
•Fortnightly  Review,'  Nov.  1867;  April.  1,  1868;  July  1,  1869,  sine©  sep- 
arately published. 


128  THE  DESCENT  OF  MAN. 

the  first  impulse  to  benevolent  actions.  Habits,  moreover,  fol- 
lowed during  many  generations  probably  tend  to  be  inherited. 

But  another  and  much  more  powerful  stimulus  to  the  develop- 
ment of  the  social  virtues,  is  afforded  by  the  praise  and  the  blame 
of  our  fellow-men.  To  the  instinct  of  sympathy,  as  we  have  al- 
ready seen,  it  is  primarily  due,  that  we  habitually  bestow  both 
praise  and  blame  on  others,  whilst  we  love  the  former  and  dread 
the  latter  when  applied  to  ourselves;  and  this  instinct  no  doubt 
was  originally  acquired,  like  all  the  other  social  instincts,  through 
natural  selection.  At  how  early  a  period  the  progenitors  of  man 
in  the  course  of  their  development,  became  capable  of  feeling  and 
being  impelled  by,  the  praise  or  blame  of  their  fellow-creatures, 
we  cannot  of  course  say.  But  it  appears  that  even  dogs  appre- 
ciate encouragement,  praise,  and  blame.  The  rudest  savages  feel 
the  sentiment  of  glory,  as  they  clearly  show  by  preserving  the 
trophies  of  their  prowess,  by  their  habit  of  excessive  boasting, 
and  even  by  the  extreme  care  v/hich  they  take  of  their  personal 
appearance  and  decorations;  for  unless  they  regarded  the  opinion 
of  their  comrades,  such  habits  would  be  senseless. 

They  certainly  feel  shame  at  the  breach  of  some  of  their  lesser 
rules,  and  apparently  remorse,  as  shown  by  the  case  of  the  Aus- 
tralian who  grew  thin  and  could  not  rest  from  having  delayed 
to  murder  some  other  woman,  so  as  to  propitiate  his  dead  wife's 
spirit.  Though  I  have  not  met  with  any  other  recorded  case, 
it  is  scarcely  credible  that  a  savage,  who  will  sacrifice  his  life 
rather  than  betray  his  tribe,  or  one  who  will  deliver  himself  up 
as  a  prisoner  rather  than  break  his  parole,^  would  not  feel  re- 
morse in  his  inmost  soul,  if  he  had  failed  in  a  duty,  which  he 
held  sacred. 

f~  We  may  therefore  conclude  that  primeval  man,  at  a  very  remote 
period,  was  influenced  by  the  praise  and  blame  of  his  fellows. 
It  is  obvious,  that  the  members  of  the  same  tribe  would  approve 
of  conduct  which  appeared  to  them  to  be  for  the  general  good, 
and  would  reprobate  that  which  appeared  evil.  To  do  good  unto 
others — to  do  unto  others  as  ye  would  they  should  do  unto  you — 
is  the  foundation-stone  of  morality.  It  is,  therefore,  hardly  pos- 
sible to  exaggerate  the  importance  during  rude  times  of  the  love 
of  praise  and  the  dread  of  blame.  A  man  who  was  not  impelled 
by  any  deep,  instinctive  feeling,  to  sacrifice  his  life  for  the  good 
of  others,  yet  was  roused  to  such  actions  by  a  sense  of  glory, 
would  by  his  example  excite  the  same  wish  for  glory  in  other 
men,  and  would  strengthen  by  exercise  the  noble  feeling  of  ad- 
miration.    He  might  thus  do  far  more  good  to  his  tribe  than 

•  Mr.  Wallace  gives  cases  in  his  'Contributions  to  the  Theory  of 
Natural  Selection,'  1870,  p.  364. 


MORAL   FACULTIES.  129 

by  Degetting  offspring  with  a  tendency  to  inherit  his  own  high 
character. 

With  increased  experience  and  reason,  man  perceives  the  more  ) 
remote  consequences  of  his  actions,  and  the  self-regarding  vir-  / 
tues,  such  as  temperance,  chastity,  &c.,  which  during  early  times/ 
are,  as  we  have  before  seen,  utterly   disregarded,   come  to  be 
highly  esteemed  or  even  held  sacred.    I  need  not,  however,  repeat 
what  I  have  said  on  this  head  in  the  fourth  chapter.    Ultimately 
our  moral  sense  or  conscience  becomes  a  highly  complex  senti- 
ment— originating  in  the  social  instincts,  largely  guided  by  the 
approbation  of  our  fellow-men,  ruled  by  reason,  self-interest,  and 
in  later  times  by  deep  religious  feelings,  and  confirmed  by  instruc- 
tion and  habit. 

It  must  not  be  forgotten  that  although  a  high  standard  of 
morality  gives  but  a  slight  or  no  advantage  to  each  individual 
man  and  his  children  over  the  other  men  of  the  same  tribe,  yet 
that  an  increase  in  the  number  of  well-endowed  men  and  an 
advancement  in  the  standard  of  morality  will  certainly  give  an 
immense  advantage  to  one  tribe  over  another.  A  tribe  including  - 
many  members  who,  from  possessing  in  a  high  degree  the  spirit 
of  patriotism,  fidelity,  obedience,  courage,  and  sympathy,  were 
always  ready  to  aid  one  another,  and  to  sacrifice  themselves  for 
the  common  good,  would  be  victorious  over  most  other  tribes; 
and  this  would  be  natural  selection.  At  all  times  throughout 
the  world  tribes  have  supplanted  other  tribes;  and  as  morality 
is  one  important  element  in  their  success,  the  standard  of 
morality  and  the  number  of  well-endowed  men  will  thus  every- 
where tend  to  rise  and  increase.  .^ 

It  is  however,  very  difficult  to  form  any  judgment  why  one 
particular  tribe  and  not  another  has  been  successful  and  has 
risen  in  the  scale  of  civilization.  Many  savages  are  in  the  same 
condition  as  when  first  discovered  several  centuries  ago.  As  Mr. 
Bagehot  has  remarked,  we  are  apt  to  look  at  progress  as  normal 
in  human  society;  but  history  refutes  this.  The  ancients  did 
not  even  entertain  the  idea,  nor  do  the  Oriental  nations  at  the 
present  day.  According  to  another  high  authority.  Sir  Henry 
Maine,^  "the  greatest  part  of  mankind  has  never  shown  a  particle 
"of  desire  that  its  civil  institutions  should  be  improved."  Progress 
seems  to  depend  on  many  concurrent  favorable  conditions,  far 
too  complex  to  be  followed  out.  But  it  has  often  been  remarked, 
that  a  cool  climate,  from  leading  to  industry  and  to  the  various 
arts,  has  been  highly  favorable  thereto.  The  Esquimaux,  pressed 
by  hard  necessity,  have  succeeded  in  many  ingenious  inventions, 
but  their  climate  has  been  too  severe  for  continued  progress. 

'  'Ancient  Law,'  1861,  p.  22.    For  Mr.  Bagehot's  remarks,  'Fortnightly 
Review,'  April  1,  1368,  p.  452. 
10 


130  THE   DESCENT  OF  MAN. 

Nomadic  habits,  whether  over  wide  plains,  or  through  the  dense 
forests  of  the  tropics,  or  along  the  shores  of  the  sea,  have  in 
every  case  been  highly  detrimental.  Whilst  observing  the  bar- 
barous inhabitants  of  Tierra  del  Puego,  it  struck  me  that  the  pos- 
session of  some  property,  a  fixed  abode,  and  the  union  of  many 
families  under  a  chief,  were  the  indispensable  requisites  for  civil- 
ization. Such  habits  almost  necessitate  the  cultivation  of  the 
ground;  and  the  first  steps  in  cultivation  would  probably  result, 
as  I  have  elsewhere  shown,^  from  some  such  accident  as  the  seeds 
of  a  fruit-tree  falling  on  a  heap  of  refuse,  and  producing  an  un- 
usually fine  variety.  The  problem,  however,  of  the  first  advance 
of  savages  towards  civilization  is  at  present  much  too  difficult  to 
be  solved. 

Natural  Selection  as  affecting  Civilized  Nations. — I  have  hither- 
to only  considered  the  advancement  of  man  from  a  semi-human 
condition  to  that  of  the  modern  savage.  But  some  remarks  on 
the  action  of  natural  selection  on  civilized  nations  may  be  worth 
adding.  This  subject  has  been  ably  discussed  by  Mr.  W.  R. 
Greg,^  and  previously  by  Mr.  Wallace  and  Mr.  Galton."  Most 
of  my  remarks  are  taken  from  these  three  authors.  With  sav- 
ages, the  weak  in  body  or  mind  are  soon  eliminated;  and  those 
that  survive  commonly  exhibit  a  vigorous  state  of  health.  We 
Civilized  men,  on  the  other  hand,  do  our  utmost  to  check  the 
process  of  elimination;  we  build  asylums  for  the  imbecile,  the 
maimed,  and  the  sick;  we  institute  poor-laws;  and  our  medical 
men  exert  their  utmost  skill  to  save  the  life  of  every  one  to  the 
last  moment.  There  is  reason  to  believe  that  vaccination  has 
preserved  thousands,  who  from  a  weak  constitution  would  for- 
merly have  succumbed  to  small-pox.  Thus  the  weak  members  of 
civilized  societies  propagate  their  kind.  No  one  who  has  attended 
to  the  breeding  of  domestic  animals  will  doubt  that  this  must 
be  highly  injurious  to  the  race  of  man.  It  is  surprising  how  soon 
a  want  of  care,  or  care  wrongly  directed,  leads  to  the  degenera- 


8  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  i. 
p.  309. 

»  'Fraser's  Mag-azine,'  Sept.  1868,  p.  353.  This  article  seems  to  have 
struck  many  persons,  and  has  given  rise  to  two  remarkable  essays, 
and  a  rejoinder  in  the  'Spectator,'  Oct.  3rd  and  17th,  1868.  It  has  also 
been  discussed  in  the  'Q.  Journal  of  Science,'  1869,  p.  152,  and  by  Mr. 
JLiawson  Tait  in  the  'Dublin  Q.  Journal  of  Medical  Science,'  Feb.  1S69, 
and  by  Mr.  E.  Ray  Lankester  in  his  'Comparative  Longevity,'  1870,  p. 
128.  Similar  views  appeared  previously  in  tHe  'Australasian,'  July  13, 
1867.    I  have  borrowed  ideas  from  several  of  these  writers. 

10  For  Mr.  Wallace,  see  'Anthropolog.  Review,'  as  before  cited.  Mr. 
Galton  in  'MacmiUan's  Magazine,'  Aug.  1865.  p;  318;  also  his  great  work, 
'Hereditary  Genius,'  1870. 


CIVILIZED   NATIONS.  131 

tlon  of  a  domestic  race;  but  excepting  in  the  case  of  man  him- 
self, hardly  any  one  is  so  ignorant  as  to  allow  his  worst  animals 
to  breed. 

The  aid  which  we  feel  impelled  to  give  to  the  helpless  is  mainly 
an  incidental  result  of  the  instinct  of  sympathy,  which  was  orig- 
inally acquired  as  part  of  the  social  instincts,  but  subsequently 
rendered,  in  the  manner  previously  indicated,  more  tender  and 
more  widely  diffused.  Nor  could  we  check  our  sympathy,  even 
at  the  urging  of  hard  reason,  without  deterioration  in  the  noblest 
part  of  our  nature.  The  surgeon  may  harden  himself  whilst  per- 
forming an  operation,  for  he  knows  that  he  is  acting  for  the  good 
of  his  patient;  but  if  we  were  intentionally  to  neglect  the  weak  and 
helpless,  it  could  only  be  for  a  contingent  benefit,  with  an  over- 
whelming present  evil.  We  must  therefore  bear  the  undoubtedly 
bad  effects  of  the  weak  surviving  and  propagating  their  kind;  but 
there  appears  to  be  at  least  one  check  in  steady  action,  namely 
that  the  weaker  and  inferior  members  of  society  do  not  marry 
so  freely  as  the  sound;  and  this  check  might  be  indefinitely  in- 
creased by  the  weak  in  body  or  mind  refraining  from  marriage, 
though  this  is  more  to  be  hoped  for  than  expected. 

In  every  country  in  which  a  large  standing  army  is  kept  up, 
the  finest  young  men  are  taken  by  the  conscription  or  are  en- 
listed. They  are  thus  exposed  to  early  death  during  war,  are 
often  tempted  into  vice,  and  are  prevented  from  marrying  during 
the  prime  of  life.  On  the  other  hand  the  shorter  and  feebler  men, 
with  poor  constitutions,  are  left  at  home,  and  consequently  have 
a  much  better  chance  of  marrying  and  propagating  their  kind." 

Man  accumulates  property  and  bequeaths  it  to  his  children, 
so  that  the  children  of  the  rich  have  an  advantage  over  the  poor 
in  the  race  for  success,  independently  of  bodily  or  mental  su- 
periority. On  the  other  hand,  the  children  of  parents  who  are 
short-lived,  and  are  therefore  on  an  average  deficient  in  health 
and  vigor,  come  into  their  property  sooner  than  other  children, 
and  will  be  likely  to  marry  earlier,  and  leave  a  larger  number  of 
offspring  to  inherit  their  inferior  constitutions.  But  the  in- 
heritance of  property  by  itself  is  very  far  from  an  evil;  for  with- 
out the  accumulation  of  capital  the  arts  could  not  progress;  and 
it  is  chiefly  through  their  power  that  the  civilized  races  have 
extended,  and  are  now  everywhere  extending  their  range,  so  as 
to  take  the  place  of  the  lower  races.  Nor  does  the  moderate  ac- 
cumulation of  wealth  interfere  with  the  process  of  selection. 
When  a  poor  man  becomes  moderately  rich,  his  children  enter 
trades  or  professions  in  which  there  is  struggle  enough,  so  that 


"  Prof.  H.  Fick  ('Einfluss  der  Naturwissenschaft  auf  das  Recht,' 
June  1872,)  has  some  good  remarks  on  this  head,  and  on  other  such 
points. 


132  f  HE3  DESCENT  OF  MAN. 

the  able  in  body  and  mind  succeed  best.  The  presence  of  a  body 
of  well-instructed  men,  who  have  not  to  labor  for  their  daily 
bread,  is  important  to  a  degree  which  cannot  be  over-estimated; 
as  all  high  intellectual  work  is  carried  on  by  them,  and  on  such 
work,  material  progress  of  all  kinds  mainly  depends,  not  to 
mention  other  and  higher  advantages.  No  doubt  wealth  when 
very  great  tends  to  convert  men  into  useless  drones,  but  their 
number  is  never  large;  and  some  degree  of  elimination  here 
occurs,  for  we  daily  see  rich  men,  who  happen  to  be  fools  or 
profligate,  squandering  away  their  wealth. 

Primogeniture  with  entailed  estates  is  a  more  direct  evil, 
though  it  may  formerly  have  been  a  great  advantage  by  the 
creation  of  a  dominant  class,  and  any  government  is  better  than 
none.  Most  eldest  sons,  though  they  may  be  weak  in  body  or 
mind,  marry,  whilst  the  younger  sons,  however  superior  in  these 
respects,  do  not  so  generally  marry.  Nor  can  worthless  eldest  sons 
with  entailed  estates  squander  their  wealth.  But  here,  as  else- 
where, the  relations  of  civilized  life  are  so  complex  that  some 
compensatory  checks  intervene.  The  men  who  are  rich  through 
primogeniture  are  able  to  select  generation  after  generation  the 
more  beautiful  and  charming  women;  and  these  must  generally 
be  healthy  in  body  and  active  in  mind.  The  evil  consequences, 
such  as  they  may  be,  of  the  continued  preservation  of  the  same 
line  of  descent,  without  any  selection,  are  checked  by  men  of 
rank  always  wishing  to  increase  their  wealth  and  power;  and 
this  they  effect  by  marrying  heiresses.  But  the  daughters  of 
parents  who  have  produced  single  children,  are  themselves,  as 
Mr.  Galton^'  has  shown,  apt  to  be  sterile;  and  thus  noble  families 
are  continually  cut  off  in  the  direct  line,  and  their  wealth  flows 
into  some  side  channel;  but  unfortunately  this  channel  is  not 
determined  by  superiority  of  any  kind. 

Although  civilization  thus  checks  in  many  ways  the  action  of 
natural  selection,  it  apparently  favors  the  better  development 
of  the  body,  by  means  of  good  food  and  the  freedom  from  occa- 
sional hardships.  This  may  be  inferred  from  civilized  men  hav- 
ing been  found,  wherever  compared,  to  be  physically  stronger 
than  savages.'^^  They  appear  also  to  have  equal  powers  of  en- 
durance, as  has  been  proved  in  many  adventurous  expeditions. 
Even  the  great  luxury  of  the  rich  can  be  but  little  detrimental, 
for  the  expectation  of  life  of  our  aristocracy,  at  all  ages  and  of 
both  sexes,  is  very  little  inferior  to  that  of  healthy  English  lives 
in  the  lower  classes." 


12  'Hereditary  Genius,'  1870,  pp.  132-140. 

13  Quatrefages,  'Revue  des  Cours  Scientifiques,'  1867-68,  p.  659. 

1*  See  the  fifth  and  sixth  columns,  compiled  from  good  authorities,  in 
the  table  given  in  Mr.  E,  E.  Lankester's  'Comparative  Longevity,'  1870, 

p.  115. 


CIVILIZED    NATIONS.  133 

We  will  now  look  to  the  intellectual  faculties.  If  in  each  grade 
of  society  the  members  were  divided  into  two  equal  bodies,  the 
one  including  the  intellectually  superior  and  the  other  the  in- 
ferior, there  can  be  little  doubt  that  the  former  would  succeed 
best  in  all  occupations,  and  rear  a  greater  number  of  children. 
Even  in  the  lowest  walks  of  life,  skill  and  ability  must  be  of 
some  advantage;  though  in  many  occupations,  owing  to  the  great 
division  of  labor,  a  very  small  one.  Hence  in  civilized  nations 
there  will  be  some  tendency  to  an  increase  both  in  the  number 
and  in  the  standard  of  the  intellectually  able.  But  I  do  not  wish 
to  assert  that  this  tendency  may  not  be  more  than  counterbal- 
anced in  other  ways  as  by  the  multiplication  of  the  reckless  and 
improvident;  but  even  to  such  as  these,  ability  must  be  some 
advantage. 

It  has  often  been  objected  to  views  like  the  foregoing,  that  the 
most  eminent  men  who  have  ever  lived  have  left  no  offspring  to 
inherit  their  great  intellect.  Mr.  Galton  says,^^  "I  regret  I  am 
"unable  to  solve  the  simple  question  whether,  and  how  far,  men 
"and  women  who  are  prodigies  of  genius  are  infertile.  I  have, 
"however,  shown  that  men  of  eminence  are  by  no  means  so." 
Great  lawgivers,  the  founders  of  beneficent  religions,  great  philos- 
ophers and  discoverers  in  science,  aid  the  progress  of  mankind 
in  a  far  higher  degree  by  their  works  than  by  leaving  a  numerous 
progeny.  In  the  case  of  corporeal  structures,  it  is  the  selection  of 
the  slightly  better-endowed  and  the  elimination  of  the  slightly 
less  well-endowed  individuals,  and  not  the  preservation  of 
strongly-marked  and  rare  anomalies,  that  leads  to  the  advance- 
ment of  a  species.^*^  So  it  will  be  with  the  intellectual  faculties, 
since  the  somewhat  abler  men  in  each  grade  of  society  succeed 
rather  better  than  the  less  able,  and  consequently  increase  in 
number,  if  not  otherwise  prevented.  When  in  any  nation  the 
standard  of  intellect  and  the  number  of  intellectual  men  have 
increased,  we  may  expect  from  the  law  of  the  deviation  from 
an  average,  that  prodigies  of  genius  will,  as  shown  by  Mr.  Galton, 
appear  somewhat  more  frequently  than  before. 

In  regard  to  the  moral  qualities,  some  elimination  of  the  worst 
dispositions  is  always  in  progress  even  in  the  most  civilized  na- 
tions. Malefactors  are  executed,  or  imprisoned  for  long  periods, 
so  that  they  cannot  freely  transmit  their  bad  qualities.  Melan- 
cholic and  insane  persons  are  confined,  or  commit  suicide.  Violent 
and  quarrelsome  men  often  come  to  a  bloody  end.  The  restless 
who  will  not  follow  any  steady  occupation — and  t^iis  relic  of 
barbarism  is  a  great  check  to  civilization^' — emigrate  to  newly- 

15  'Hereditary  Genius,'  1870,  p.  330. 

"  'Origin  of  Species'  (fifth  edition,  1869),  p.  104. 

"  'Hereditar:--  Genius,'  1870,  p.  347. 

10 


134  THE  DESCENT  OF  MAN. 

settled  countries,  where  they  prove  useful  pioneers.  Intemper- 
ance is  so  highly  destructive,  that  the  expectation  of  life  of  the 
intemperate,  at  the  age  of  thirty  for  instance,  is  only  13.8  years; 
whilst  for  the  rural  laborers  of  England  at  the  same  age  it  is 
40.59  years.^^  Profligate  women  bear  few  children,  and  profligate 
men  rarely  marry;  both  suffer  from  disease.  In  the  breeding  of 
domestic  animals,  the  elimination  of  those  individuals,  though 
few  in  number,  which  are  in  any  marked  manner  inferior,  is  by 
no  means  an  unimportant  element  towards  success.  This  espe- 
cially holds  good  with  injurious  characters  which  tend  to  reap- 
pear through  reversion,  such  as  blackness  in  sheep;  and  with 
mankind  some  of  the  worst  dispositions,  which  occasionally  with- 
out any  assignable  cause  make  their  appearance  in  families,  may 
perhaps  be  reversions  to  a  savage  state,  from  which  we  are  not 
removed  by  very  many  generations.  This  view  seems  indeed 
recognized  in  the  common  expression  that  such  men  are  the  black 
sheep  of  the  family. 

With  civilized  nations,  as  far  as  an  advanced  standard  of 
morality,  and  an  increased  number  of  fairly  good  men  are  con- 
cerned, natural  selection  apparently  effects  but  little;  though 
the  fundamental  social  instincts  were  originally  thus  gained. 
But  I  have  already  said  enough,  whilst  treating  of  the  lower 
races,  on  the  causes  which  lead  to  the  advance  of  morality,  name- 
ly, the  approbation  of  our  fellow-men — ^the  strengthening  of  our 
sympathies  by  habit — example  and  imitation — reason — experience, 
and  even  self-interest — instruction  during  youth,  and  religious 
feelings. 

A  most  important  obstacle  in  civilized  countries  to  an  increase 
in  the  number  of  men  of  a  superior  class  has  been  strongly  in- 
sisted on  by  Mr.  Greg  and  Mr.  Galton,^^  namely,  the  fact  that  the 
very  poor  and  reckless,  who  are  often  degraded  by  vice,  almost 
invariably  marry  early,  whilst  the  careful  and  frugal,  who  are 
generally  otherwise  virtuous,  marry  late  in  life,  so  that  they  may 
be  able  to  support  themselves  and  their  children  in  comfort. 
Those  who  marry  early  produce  within  a  given  period  not  only  a 
greater  number  of  generations,  but,  as  shown  by  Dr.  Duncan,^" 


1*  E.  Ray  Lankester,  'Comparative  Longevity,'  1870,  p.  115.  The  table 
of  the  intemperate  is  from  Nelson's  'Vital  Statistics.'  In  regard  to 
profligacy,  see  Dr.  Farr,  'Influence  of  Marriage  on  Mortality/  'Nat. 
Assoc,  for  the  Promotion  of  Social  Science,'  1858. 

i»  'Fraser's  Magazine,'  Sept.  1868,  p.  353.  'Macmillan's  Magazine,'  Aug 
1865,  p.  318.  The  Rev.  F.  W.  Farrar  ('Fraser's  Mag.'  Aug.  1870,  p.  264), 
takes  a  different  view. 

20  'On  the  Laws  of  the  Fertility  of  Women,'  in  'Transact.  Royal 
Soc'  Edinburgh,  vol.  xxiv.  p.  287;  now  published  separately  under  the 
title  of  'Fecundity,  Fertility,  and  Sterility,'  1871.  See,  also,  Mr.  Galtoiis- 
'Hereditary  Genius,'  pp.  352-357,  fot  observations  to  the  above  effect. 


CIVILIZED   NATIONS.  135 

they  produce  many  more  children.  The  children,  moreover,  that 
are  born  by  mothers  during  the  prime  of  life  are  heavier  and 
larger,  and  therefore  probably  more  vigorous,  than  those  born 
at  other  periods.  Thus  the  reckless,  degraded,  and  often  vicious 
members  of  society,  tend  to  increase  at  a  quicker  rate  than  the 
provident  and  geiuerally  virtuous  members.  Or  as  Mr.  Greg  puts 
the  case:  "The  careless,  squalid,  unaspiring  Irishman  multiplies 
"like  rabbits:  the  frugal,  foreseeing,  self-respecting,  ambitious 
"Scot,  stern  in  his  morality,  spiritual  in  his  faith,  sagacious  and 
"disciplined  in  his  intelligence,  passes  his  best  years  in  struggle 
"and  in  celibacy,  marries  late,  and  leaves  few  behind  him.  Given 
"a  land  originally  peopled  by  a  thousand  Saxons  and  a  thousand 
"Celts — and  in  a  dozen  generations  five-sixths  of  the  population 
"would  be  Celts,  but  five-sixths  of  the  property,  of  the  power,  of 
"the  intellect,  would  belong  to  the  one-sixth  of  Saxons  that  re- 
"mained.  In  the  eternal  'struggle  for  existence,'  it  would  be  the 
"inferior  and  less  favored  race  that  had  prevailed — and  prevailed 
"by  virtue  not  of  its  good  qualities  but  of  its  faults." 

There  are,  however,  some  checks  to  this  downward  tendency. 
We  have  seen  that  the  intemperate  suffer  from  a  high  rate  of 
mortality,  and  the  extremely  profligate  leave  few  offspring.  The 
poorest  classes  crowd  into  towns,  and  it  has  been  proved  by  Dr. 
Stark  from  the  statistics  of  ten  years  in  Scotland,-^  that  at  all 
ages  the  death-rate  is  higher  in  towns  than  in  rural  districts, 
"and  during  the  first  five  years  of  life  the  town  death-rate  is 
"almost  exactly  double  that  of  the  rural  districts."  As  these  re- 
turns include  both  the  rich  and  the  poor,  no  doubt  more  than 
twice  the  number  of  births  would  be  requisite  to  keep  up  the 
number  of  the  very  poor  inhabitants  in  the  towns,  relatively  to 
those  in  the  country.  With  women,  marriage  at  too  early  an 
age  is  highly  injurious;  for  it  has  been  found  in  France  that, 
"twice  as  many  wives  under  twenty  die  in  the  year,  as  died  out 
"of  the  same  number  of  the  unmarried."  The  mortality,  also, 
of  husbands  under  twenty  is  "excessively  high,"^^  but  what  the 
cause  of  this  may  be,  seems  doubtful.  Lastly,  if  the  men  who 
prudently  delay  marrying  until  they  can  bring  up  their  families 
in  comfort,  were  to  select,  as  they  often  do,  women  in  the  prime 
of  life,  the  rate  of  increase  in  the  better  class  would  be  only 
slightly  lessened. 

It  was  established  from  an  enormous  body  of  statistics,  taken 
during  1853,  that  the  unmarried  men  throughout  France,  between 

21  'Tenth  Annual  Report  of  Births,  Deaths,  &c.,  in  Scotland,'  1867,  p. 
xxix. 

22  These  quotations  are  taken  from  our  highest  authority  on  such 
questions,  namely.  Dr.  Farr,  in  his  paper  'On  the  Influence  of  Mar- 
riage on  the  Mortality  of  the  French  People,'  read  before  the  Nat. 
Assoc,  for  the  Promotion  of  Social  Science  1858. 


136  THE  DESCENT  OP  MAN. 

the  ages  of  twenty  and  eighty,  die  in  a  much  larger  proportion 
than  the  married:  for  instance,  out  of  every  1000  unmarried  men, 
between  the  ages  of  twenty  and  thirty,  11.3  annually  died,  whilst 
of  the  married  only  6.5  died.^^  A  similar  law  was  proved  to  hold 
good,  during  the  years  1863  and  1864,  with  the  entire  population 
above  the  age  of  twenty  in  Scotland:  for  instance,  out  of  every 
1000  unmarried  men,  between  the  ages  of  twenty  and  thirty,  14.97 
annually  died,  whilst  of  the  married  only  7.24  died,  that  is  less 
than  half.-*  Dr.  Stark  remarks  on  this,  "Bachelorhood  is  more 
"destructive  to  life  than  the  most  unwholesome  trades,  or  than 
"residence  in  an  unwholesome  house  or  district  where  there  has 
"never  been  the  most  distant  attempt  at  sanitary  improvement." 
He  considers  that  the  lessened  mortality  is  the  direct  result  of 
"marriage,  and  the  more  regular  domestic  habits  which  attend 
"that  state."  He  admits,  however,  that  the  intemperate,  profli- 
gate, and  criminal  classes,  whose  duration  of  life  is  low,  do  not 
commonly  marry;  and  it  must  likewise  be  admitted  that  men 
with  a  weak  constitution,  ill  health,  or  any  great  infirmity  in  body 
or  mind,  will  often  not  wish  to  marry,  or  will  be  rejected.  Dr. 
Stark  seems  to  have  come  to  the  conclusion  that  marriage  in  itself 
is  a  main  cause  of  prolonged  life,  from  finding  that  aged  married 
men  still  have  a  considerable  advantage  in  this  respect  over  the 
unmarried  of  the  same  advanced  age;  but  every  one  must  have 
known  instances  of  men,  who  with  weak  health  during  youth 
did  not  marry,  and  yet  have  survived  to  old  age,  though  remaining 
weak,  and  therefore  always  with  a  lessened  chance  of  life  or  of 
marrying.  There  is  another  remarkable  circumstance  which 
seems  to  support  Dr.  Stark's  conclusion,  namely,  that  widows  and 
widowers  in  France  suffer  in  comparison  with  the  married  a  very 
heavy  rate  of  mortality;  but  Dr.  Farr  attributes  this  to  the  pov- 
erty and  evil  habits  consequent  on  the  disruption  of  the  family, 
and  to  grief.  On  the  whole  we  may  conclude  with  Dr.  Farr  that 
the  lesser  mortality  of  married  than  of  unmarried  men,  which 
seems  to  be  a  general  law,  "is  mainly  due  to  the  constant  elimina- 
"tion  of  imperfect  types,  and  to  the  skillful  selection  of  the  finest 
"individuals  out  of  each  successive  generation;"  the  selection  re- 
lating only  to  the  marriage  state,  and  acting  on  all  corporeal,  in- 
tellectual, and  moral  qualities.^^    We  may,  therefore,  infer  that 

23  Dr.  Farr,  ibid.  The  quotations  given  below  are  extracted  from  the 
same  striliing  paper. 

24  I  have  taken  the  mean  of  the  quinquennial  means,  given  in  'The 
Tenth  Annual  Report  of  Births,  Deaths,  &c.,  in  Scotland,'  1867.  The 
quotation  from  Dr.  Stark  is  copied  from  an  article  in  the  'Daily  News,' 
Oct.  17th,  1S6S,  which  Dr.  Farr  considers  very  carefully  written. 

25  Dr.  Duncan  remarks  ('Fecundity,  Fertility,'  &c.,  1871,  p.  334)  on  this 
subject:  "At  every  age  the  healthy  and  beautiful  go  over  from  the 
"unmarried  side  to  the  married,  leaving  the  unmarried  columns 
"crowded  with  the  sickly  and  unfortunate." 


CIVILIZED    NATIONS.  137 

sound  and  good  men  who  out  of  prudence  remain  for  a  time  un- 
married, do  not  suffer  a  high  rate  of  mortality. 

If  the  various  checks  specified  in  the  two  last  paragraphs,  and 
perhaps  others  as  yet  unknown,  do  not  prevent  the  reckless,  the  | 
vicious  and  otherwise  inferior  members  of  society  from  increas-  i 
ing  at  a  quicker  rate  than  the  better  class  of  men,  the  nation  will 
retrograde,  as  has  too  often  occurred  in  the  history  of  the  world. 
We  must  remember  that  progress  is  no  invariable  rule.  It  is 
very  difficult  to  say  why  one  civilized  nation  rises,  becomes  more 
powerful,  and  spreads  more  widely,  than  another;  or  why  the 
same  nation  progresses  more  quickly  at  one  time  than  at  another. 
We  can  only  say  that  it  depends  on  an  increase  in  the  actual  num- 
ber of  the  population  on  the  number  of  the  men  endowed  with 
high  intellectual  and  moral  faculties,  as  well  as  on  their  standard 
of  excellence.  Corporeal  structure  appears  to  have  little  influence, 
except  so  far  as  vigor  of  body  leads  to  vigor  of  mind. 

It  has  been  urged  by  several  writers  that  as  high  intellectual 
powers  are  advantageous  to  a  nation,  the  old  Greeks,  who  stood 
some  grades  higher  in   intellect  than   any  race  that  has   ever 
existed,'^  ought,  if  the  power  of  natural  selection  were  real,  to 
have  risen  still  higher  in  the  scale,  increased  in  number,  and 
stocked  the  whole  of  Europe.    Here  we  have  the  tacit  assumption, 
so  often  made  with  respect  to  corporeal  structures,  that  there  is 
some  innate  tendency  towards  continued  development  in  mind  and  , 
body.    But  development  of  all  kinds  depends  on  many  concur-  \ 
rent  favorable  circumstances.    Natural  selection  acts  only  tenta-  t 
tively.    Individuals  and  races  may  have  acquired  certain  indis- 
putable advantages,  and  yet  have  perished  from  failing  in  other     /^     . 
characters.    The  Greeks  may  have  retrograded  from  a  want  of     ^^r-^' 
coherence  between  the  many  small  states,  from  the  small  size  of 
their  whole  country,  from  the  practice  of  slavery,  or  from  extreme 
sensuality;  for  they  did  not  succumb  until  "they  were  enervated 
"and  corrupt  to  the  very  core."-^    The  western  nations  of  Europe, 
who  now  so  immeasurably  surpass  their  former  savage  progeni- 
tors, and  stand  at  the  summit  of  civilization,  owe  little  or  none 
of  their  superiority  to  direct  inheritance  from  the  old  Greeks, 
though  they  owe  much  to  the  written  works  of  that  wonderful 
people. 

Who  can  positively  say  why  the  Spanish  nation,  so  dominant  at 
one  time,  has  been  distanced  in  the  race.  The  awakening  of  the 
nations  of  Europe  from  the  dark  ages  is  a  still  more  perplexing 
problem.  At  that  early  period,  as  Mr.  Galton  has  remarked,  al- 
most all  the  men  of  a  gentle  nature,  those  given  to  meditation  or 

2^  See  the  ingenious  and  original  argument  on  this  subject  by  Mr.  Gal- 
ton, 'Hereditary  Genius,'  pp.  340-342. 
27  Mr.  Greg  'Fraser's  Magazine,'  Sept.  1868,  p.  357. 


138  THE  DESCENT  OF  MAN. 

culture  of  the  mind,  had  no  refuge  except  in  the  bosom  of  a 
Church  which  demanded  celibacy;-^  and  this  could  hardly  fail  to 
have  had  a  deteriorating  influence  on  each  successive  generation. 
During  this  same  period  the  Holy  Inquisition  selected  with  ex- 
treme care  the  freest  and  boldest  men  in  order  to  burn  or  im- 
prison them.  In  Spain  alone  some  of  the  best  men — those  who 
doubted  and  questioned,  and  without  doubting  there  can  be  no 
progress — were  eliminated  during  three  centuries  at  the  rate  of  a 
thousand  a  year.  The  evil  which  the  Catholic  Church  has  thus 
effected  is  incalculable,  though  no  doubt  counterbalanced  to  a  cer- 
tain, perhaps  to  a  large,  extent  in  other  ways;  nevertheless, 
Europe  has  progressed  at  an  unparalleled  rate. 
"  The  remarkable  success  of  the  English  as  colonists,  compared 
A'to  other  European  nations,  has  been  ascribed  to  their  "daring 
"and  persistent  energy;"  a  result  which  is  well  illustrated  by  com- 
paring the  progress  of  the  Canadians  of  English  and  French  ex- 
traction; but  who  can  say  how  the  English  gained  their  energy? 
There  is  apparently  much  truth  in  the  belief  that  the  wonderful 
progress  of  the  United  States,  as  well  as  the  character  of  the  peo- 
ple, are  the  results  of  natural  selection;  for  the  more  energetic, 
restless,  and  courageous  men  from  all  parts  of  Europe  have 
emigrated  during  the  last  ten  or  twelve  generations  to  that  great 
country,  and  have  there  succeeded  best.-*  Looking  to  the  dis- 
tant future,  I  do  not  think  that  the  Rev.  Mr.  Zincke  takes  an 
exaggerated  view  when  he  says:^''  "All  other  series  of  events — 
"as  that  v/hich  resulted  in  the  culture  of  mind  in  Greece,  and 
"that  which  resulted  in  the  empire  of  Rome — only  appear  to 
"have  purpose  and  value  when  viewed  in  connection  with,  or 
"rather  as  subsidiary  to. . .  .the  great  stream  of  Anglo-Saxon  emi- 
"gration  to  the  west."  Obscure  as  is  the  problem  of  the  advance 
of  civilization,  we  can  at  least  see  that  a  nation  which  produced 
during  a  lengthened  period  the  greatest  number  of  highly  intel- 
lectual, energetic,  brave,  patriotic,  and  benevolent  men,  would 
generally  prevail  over  less  favored  nations. 

Natural  selection  follows  from  the  struggle  for  existence;  and 
this  from  a  rapid  rate  of  increase.  It  is  impossible  not  to  regret 
bitterly,  but  whether  wisely  is  another  question,  the  rate  at  which 
man  tends  to  increase;  for  this  leads  in  barbarous  tribes  to  in- 

28  'Hereditary  Genius,'  1870,  pp.  357-359.  The  Rev.  F.  W.  Farrar  'Fras- 
er's  Mag.,'  Aug.  1870,  p.  257,  advances  arguments  on  the  other  side.  Sir 
C.  Lyell  had  already  ('Principles  of  Geology,'  vol.  ii.  1868,  p.  489)  in  a 
striking  passage  called  attention  to  the  evil  influence  of  the  Holy  In- 
quisition in  having,  through  selection,  lowered  the  general  standard 
of  intelligence  in  Europe. 

29  Mr.  Galton,  'Macmillan's  Magazine,'  August,  1865,  p.  325.  See,  also 
'Nature,'  'On  Darwinism  and  National  Life,'  Dec.  1869,  p.  184. 

so  'Last  Winter  in  the  United  States,'  1868,  p.  29. 


CIVILIZED   NATIONS.  139 

fanticide  and  many  other  evils,  and  in  civilized  nations  to  abject 
poverty,  celibacy,  and  to  the  late  marriages  of  the  prudent.  But 
as  man  suffers  from  the  same  physical  evils  as  the  lower  animals, 
he  has  no  right  to  expect  an  immunity  from  the  evils  consequent 
on  the  struggle  for  existence.  Had  he  not  been  subjected  during 
primeval  times  to  natural  selection,  assuredly  he  would  never 
have  attained  to  his  present  rank.  Since  we  see  in  many  parts  of 
the  world  enormous  areas  of  the  most  fertile  land  capable  of  sup- 
porting numerous  happy  homes,  but  peopled  only  by  a  few  wan- 
dering savages,  it  might  be  argued  that  the  struggle  for  existence 
had  not  been  sufficiently  severe  to  force  man  upwards  to  his  high- 
est standard.  Judging  from  all  that  we  know  of  man  and  the 
lower  animals,  there  has  always  been  sufficient  variability  in  their 
intellectual  and  moral  faculties,  for  a  steady  advance  through 
natural  selection.  No  doubt  such  advance  demands  many  favora- 
ble concurrent  circumstances;  but  it  may  be  well  doubted  wheth- 
er the  most  favorable  would  have  sufficed,  had  not  the  rate  of  in- 
crease been  rapid,  and  the  consequent  struggle  for  existence  ex- 
tremely severe.  It  even  appears  from  what  we  see,  for  instance, 
in  parts  of  S.  America,  that  a  people  which  may  be  called  civ- 
ilized, such  as  the  Spanish  settlers,  is  liable  to  become  indolent 
and  to  retrograde,  when  the  conditions  of  life  are  very  easy.  With 
highly  civilized  nations  continued  progress  depends  in  a  subordi- 
nate degree  on  natural  selection;  for  such  nations  do  not  sup- 
plant and  exterminate  one  another  as  do  savage  tribes.  Neverthe- 
less the  more  intelligent  members  within  the  same  community 
will  succeed  better  in  the  long  run  than  the  inferior,  and  leave 
a  more  numerous  progeny,  and  this  is  a  form  of  natural  selection. 
The  more  efficient  causes  of  progress  seem  to  consist  of  a  good 
education  during  youth  whilst  the  brain  is  impressible,  and  of 
a  high  standard  of  excellence,  inculcated  by  the  ablest  and  best 
men,  embodied  in  the  laws,  customs  and  traditions  of  the  nation, 
and  enforced  by  public  opinion.  It  should,  however,  be  borne  in 
mind,  that  the  enforcement  of  public  opinion  depends  on  our 
appreciation  of  the  approbation  and  disapprobation  of  others; 
and  this  appreciation  is  founded  on  our  sympathy,  which  it  can 
hardly  be  doubted  was  originally  developed  through  natural  se- 
lection as  one  of  the  most  important  elements  of  the  social  in- 
stincts.^^ 

On  the  evidence  that  all  civilized  nations  were  once  barbarous. — 
The  present  subject  has  been  treated  in  so  full  and  admirable  a 
manner  by  Sir  J.  Lubbock,^^  Mr.  Tylor,  Mr.  M'Lennan,  and  others, 

^^  I  am  much  indebted  to  Mr.  John  Morley  for  some  good  criticisms 
on  this  subject:  see,  also,  Broca,  'Les  Selections,'  'Revue  d'Anthro- 
pologie,'   1872. 

32  'On  the  Origin  of  Civilization,'  'Proc.  Ethnological  Soc'  Nov.  26, 
1867. 


140  THE   DESCENT  OP  MAN. 

that  I  need  here  give  only  the  briefest  summary  of  their  results. 
The  arguments  recently  advanced  by  the  Duke  of  Argyll'^*  and 
formerly  by  Archbishop  Whately,  in  favor  of  the  belief  that  man 
came  into  the  world  as  a  civilized  being,  and  that  all  savages 
have  since  undergone  degradation,  seem  to  me  weak  in  com- 
parison with  those  advanced  on  the  other  side.  Many  nations,  no 
doubt,  have  fallen  away  in  civilization,  and  some  may  have  lapsed 
into  utter  barbarism,  though  on  this  latter  head  I  have  met  with 
no  evidence.  The  Fuegians  were  probably  compelled  by  other 
conquering  hordes  to  settle  in  their  inhospitable  country,  and  they 
may  have  become  in  consequence  somewhat  more  degraded;  but  it 
would  be  difficult  to  prove  that  they  have  fallen  much  below  the 
Botocudos,  who  inhabit  the  finest  parts  of  Brazil. 

The  evidence  that  all  civilized  nations  are  the  descendants  of 
barbarians,  consists,  on  the  one  side,  of  clear  traces  of  their 
former  low  condition  in  still-existing  customs,  beliefs,  language. 
&c.;  and  on  the  other  side,  of  proofs  that  savages  are  independ- 
ently able  to  raise  themselves  a  few  steps  in  the  scale  of  civiliza- 
tion, and  have  actually  thus  risen.  The  evidence  on  the  first  head 
is  extremely  curious,  but  cannot  be  here  given:  I  refer  to  such 
cases  as  that  of  the  art  of  enumeration,  which,  as  Mr.  Tylor 
clearly  shows  by  reference  to  the  words  still  used  in  some  places, 
originated  in  counting  the  fingers,  first  of  one  hand  and  then  of 
the  other,  and  lastly  of  the  toes.  We  have  traces  of  this  in  our 
own  decimal  system,  and  in  the  Roman  numerals,  where,  after  the 
v.,  which  is  supposed  to  be  an  abbreviated  picture  of  a  human 
hand,  we  pass  on  to  VI.,  &c.,  when  the  other  hand  no  doubt  was 
used.  So  again,  "when  we  speak  of  three-score  and  ten,  we  are 
"counting  by  the  vigesimal  system,  each  score  thus  ideally  made, 
"standing  for  20 — for  'one  man'  as  a  Mexican  or  Carib  would  put 
"it."^*  According  to  a  large  and  increasing  school  of  philologists, 
every  language  bears  the  marks  of  its  slow  and  gradual  evolution. 
So  it  is  with  the  art  of  writing,  for  letters  are  rudiments  of  pic- 
torial representations.  It  is  hardly  possible  to  read  Mr.  M'Len- 
nan's  work^^  and  not  admit  that  almost  all  civilized  nations  still 
retain  traces  of  such  rude  habits  as  the  forcible  capture  of  wives. 
What  ancient  nation,  as  the  same  author  asks,  can  be  named 

33  'Primeval  Man,'  1869. 

3*  'Royal  Institution  of  Great  Britain,'  March  15,  1867.  Also,  'Re- 
searches into  the  Early  History  of  Mankind,'  1865. 

35  'Prihaitive  Marriage,'  1865.  See,  likewise,  an  excellent  article,  evi- 
dently by  the  same  author,  in  the  'North  British  Review,'  July,  1869. 
Also,  Mr.  L.  H.  Morgan,  'A  Conjectural  Solution  of  the  Origin  of  the 
Class  System  of  Relationship,'  in  'Proc.  American  Acad,  of  Sciences,' 
vol.  vii.  Feb.  1868.  Prof.  SchaafChausen  ('Anthropolog.  Review,'  Oct. 
1869,  p.  373)  remarks  on  "the  vestiges  of  human  sacrifices  found  both 
"in  Homer  and  the  Old  Testament." 


NATIONS.  14i 

that  was  originally  monogamous?  The  primitive  idea  of  justice, 
as  shown  by  the  law  of  battle  and  other  customs  of  which  vestiges 
still  remain,  was  likewise  most  rude.  Many  existing  superstitions 
are  the  remnants  of  former  false  religious  beliefs.  The  highest 
form  of  religion — the  grand  idea  of  God  hating  sin  and  loving 
righteousness — was  unknown  during  primeval  times. 

Turning  to  the  other  kind  of  evidence;  Sir  J.  Lubbock  has 
shown  that  some  savages  have  recently  improved  a  little  in 
some  of  their  simpler  arts.  From  the  extremely  curious  ac- 
count which  he  gives  of  the  weapons,  tools,  and  arts,  in  use 
amongst  savages  in  various  parts  of  the  world,  it  cannot  be 
doubted  that  these  have  nearly  all  been  independent  discoveries, 
excepting  perhaps  the  art  of  making  fire.'^'^  The  Australian 
boomerang  is  a  good  instance  of  one  such  independent  discovery. 
The  Tahitians  when  first  visited  had  advanced  in  many  respects 
beyond  the  inhabitants  of  most  of  the  other  Polynesian  islands. 
There  are  no  just  grounds  for  the  belief  that  the  high  culture  of 
the  native  Peruvians  and  Mexicans  was  derived  from  abroad;" 
many  native  plants  were  there  cultivated,  and  a  few  native  ani- 
mals domesticated.  We  should  bear  in  mind  that,  judging  from 
the  small  influence  of  most  missionaries,  a  wandering  crew  from 
some  semi-civilized  land,  if  washed  to  the  shores  of  America, 
would  not  have  produced  any  marked  effect  on  the  natives,  unless 
they  had  already  become  somewhat  advanced.  Looking  to  a  very 
remote  period  in  the  history  of  the  world,  we  find,  to  use  Sir  J. 
Lubbock's  well-known  terms,  a  paleolithic  and  neolithic  period; 
and  no  one  will  pretend  that  the  art  of  grinding  rough  flint  tools 
was  a  borrowed  one.  In  all  parts  of  Europe,  as  far  east  as  Greece, 
in  Palestine,  India,  Japan,  New  Zealand,  and  Africa,  including 
Egypt,  flint  tools  have  been  discovered  in  abundance;  and  of  their 
use  the^^exTsling  inhabitants  retain  no  tradition.  There  is  also 
indirect  evidence  of  their  former  use  by  the  Chinese  and  ancient 
Jews.  Hence  there  can  hardly  be  a  doubt  that  the  inhabitants  of 
these  countries,  which  include  nearly  the  whole  civilized  world, 
were  once  in  a  barbarous  condition.  To  believe  that  man  was 
aboriginally  civilized  and  then  suffered  utter  degradation  in  so 
many  regions,  is  to  take  a  pitiably  low  view  of  human  nature.  It 
is  apparently  a  truer  and  more  cheerful  view  that  progress  has 
been  much  more  general  than  retrogression;  that  man  has  risen, 
though  by  slow  and  interrupted  steps,  from  a  lowly  condition  to 
the  highest  standard  as  yet  attained  by  him  in  knowledge,  morals 
and  religion. 

38  Sir  J.  Lubbock,  'Prehistoric  Times,'  2nd  edit.  1869,  chap.  xv.  and 
xvi.  et  passim.  See,  also,  the  excellent  9th  chapter  in  Tyler's  'Early 
History  of  Mankind,'  2nd  edit.,  1870. 

3^  Dr.  F.  Muller  has  made  some  good  remarks  to  this  effect  in  the 
'Reise  der  Novara:   Anthropolog.   Theil,'   Abtheil.    iii.   1868,   s.   127. 


142  THH  DESCENT  OP  MAN. 


CHAPTER  VI. 

ON   THE  AFFINITIES   AND  GENEALOGY  OF   MAN. 

Position  of  man  in  the  animal  series — The  natural  system  genealogical 
—Adaptive  characters  of  slight  value— Various  small  points  of  re- 
semblance between  man  and  the  Quadrumana— Rank  of  man  in  the 
natural  system— Birthplace  and  antiquity  of  man— Absence  of  fossil 
connecting-links— Lower  stages  in  the  genealogy  of  man,  as  in- 
ferred, firstly  from  his  affinities  and  secondly  from  his  structured- 
Early  androgynous  condition  of  the  Vertebrata— Conclusion. 

Even  if  it  be  granted  that  the  difference  between  man  and  his 
nearest  allies  is  as  great  in  corporeal  structure  as  some  natu- 
ralists maintain,  and  although  we  must  grant  that  the  difference 
between  them  is  immense  in  mental  power,  yet  the  facts  given  in 
the  earlier  chapters  appear  to  declare,  in  the  plainest  manner, 
that  man  is  descended  from  some  lower  form,  notwithstanding 
that  connecting  links  have  not  hitherto  been  discovered. 

Man  is  liable  to  numerous,  slight,  and  diversified  variations, 
which  are  induced  by  the  same  general  causes,  are  governed 
and  transmitted  in  accordance  with  the  same  general  laws,  as  in 
the  lower  animals.  Man  has  multiplied  so  rapidly,  that  he  has 
necessarily  been  exposed  to  struggle  for  existence,  and  con- 
sequently to  natural  selection.  He  has  given  rise  to  many  races, 
some  of  which  differ  so  much  from  each  other,  that  they  have 
often  been  ranked  by  naturalists  as  distinct  species.  His  bodj"" 
is  constructed  on  the  same  homological  plan  as  that  of  other 
mammals.  He  passes  through  the  same  phases  of  embryological 
development.  He  retains  many  rudimentary  and  useless  struc- 
tures, which  no  doubt  were  once  serviceable.  Characters  occa- 
sionally make  their  re-appearance  in  him,  which  we  have  reason 
to  believe  were  possessed  by  his  early  progenitors.  If  the  origin 
of  man  has  been  wholly  different  from  that  of  all  other  animals, 
these  various  appearances  would  be  mere  empty  deceptions; 
but  such  an  admission  is  incredible.  These  appearances,  on  the 
other  hand,  are  intelligible,  at  least  to  a  large  extent,  if  man 
is  the  co-descendant  with  other  mamma,ls  of  some  unknown  and 
lower  form. 

Some  naturalists,  from  being  deeply  impressed  with  the  mental 


AFFINITIES   AND    GENEALOGY.  143 

and  spiritual  powers  of  man,  have  divided  the  whole  organic 
world  into  three  kingdoms,  the  Human,  the  Animal,  and  the 
Vegetable,  thus  giving  to  man  a  separate  kingdom.^  Spiritual 
powers  cannot  be  compared  or  classed  by  the  naturalist:  but  he 
may  endeavor  to  show,  as  I  have  done,  that  the  mental  faculties 
of  man  and  the  lower  animals  do  not  differ  in  kind,  although 
immensely  in  degree.  A  difference  in  degree,  however  great, 
does  not  justify  us  in  placing  man  in  a  distinct  kingdom,  as  will 
perhaps  be  best  illustrated  by  comparing  the  mental  powers  of 
two  insects,  namely,  a  coccus  or  scale-insect  and  an  ant,  which 
undoubtedly  belong  to  the  same  class.  The  difference  is  here 
greater  than,  though  of  a  somewhat  different  kind  from,  that 
between  man  and  the  highest  mammal.  The  female  coccus,  whilst 
young,  attaches  itself  by  its  proboscis  to  a  plant;  sucks  the  sap, 
but  never  moves  again;  is  fertilized  and  lays  eggs;  and  this  is  its 
whole  history.  On  the  other  hand,  to  describe  the  habits  and 
mental  powers  of  worker-ants,  would  require,  as  Pierre  Ruber 
has  shown,  a  large  volume;  I  may,  however,  briefly  specify  a  few 
points.  Ants  certainly  communicate  information  to  each  other, 
and  several  unite  for  the  same  work,  or  for  games  of  play.  They 
recognize  their  fellow-ants  after  months  of  absence,  and  feel  sym- 
pathy for  each  other.  They  build  great  edifices,  keep  them  clean, 
close  the  doors  in  the  evening,  and  post  sentries.  They  make 
roads  as  well  as  tunnels  under  rivers,  and  temporary  bridges  over 
them,  by  clinging  together.  They  collect  food  for  the  community, 
and  when  an  object,  too  large  for  entrance,  is  brought  to  the  nest, 
they  enlarge  the  door,  and  afterwards  build  it  up  again.  They 
store  up  seeds,  of  which  they  prevent  the  germination,  and  which, 
if  damp,  are  brought  up  to  the  surface  to  dry.  They  keep  aphides 
and  other  insects  as  milch-cows.  They  go  out  to  battle  in  regular 
bands,  and  freely  sacrifice  their  lives  for  the  common  weal.  They 
emigrate  according  to  a  preconcerted  plan.  They  capture  slaves. 
They  move  the  eggs  of  their  aphides,  as  well  as  their  own  eggs 
and  cocoons,  into  warm  parts  of  the  nest,  in  order  that  they  may 
be  quickly  hatched;  and  endless  similar  facts  could  be  given.^  On 
the  whole,  the  difference  in  mental  power  between  an  ant  and  a 
coccus  is  immense;  yet  no  one  has  ever  dreamed  of  placing  these 
insects  in  distinct  classes,  much  less  in  distinct  kingdoms.  No 
doubt  the  difference  is  bridged  over  by  other  insects;  and  this 

1  Isidore  Geoffrey  St.-Hilaire  gives  a  detailed  account  of  the  position 
assigned  to  man  bj'  various  naturalists  in  their  classifications:  'Hist. 
Nat.  Gen.'  torn.  ii.  1859,  pn.  170-189. 

2  Some  of  the  most  interesting  facts  ever  published  on  the  habits  of 
ants  are  given  by  Mr.  Belt,  in  his  'Naturalist  in  Nicaragua,'  1874.  See, 
also,  Mr.  Moggridge's  admirable  work,  'Harvesting  Ants,'  &c.,  1873, 
also  'L'Instinct  chez  les  Insectes,'  by  M.  George  Pouchet,  'Revue  des 
Deux  Mondes,'  F.eb.  1870,  p.  682. 

2 


144  THE  DESCENT  OP  MAN. 

is  not  the  case  with  man  and  the  higher  apes.  But  we  have 
every  reason  to  believe  that  the  breaks  in  the  series  are  simply 
the  results  of  many  forms  having  become  extinct. 

Professor  Owen,  relying  chiefly  on  the  structure  of  the  brain, 
has  divided  the  mammalian  series  into  four  sub-classes.  One  of 
these  he  devotes  to  man;  in  another  he  places  both  the  Marsu- 
pials and  the  Monotremata;  so  that  he  makes  man  as  distinct 
from  all  other  mammals  as  are  these  two  latter  groups  conjoined. 
This  view  has  not  been  accepted,  as  far  as  I  am  aware,  by  any 
naturalist  capable  of  forming  an  independent  judgment^  and 
therefore  need  not  here  be  further  considered. 

We  can  understand  why  a  classification  founded  on  any  single 
character  or  organ — even  an  organ  so  wonderfully  complex  and 
important  as  the  brain — or  on  the  high  development  of  the  mental 
faculties,  is  almost  sure  to  prove  unsatisfactory.  This  principle 
has  indeed  been  tried  with  hymenopterous  insects;  but  when  thus 
classed  by  their  habits  or  instincts,  the  arrangement  proved  thor- 
oughly artificial.^  Classifications  may,  of  course,  be  based  on  any 
character  whatever,  as  on  size,  color,  or  the  element  inhabited;  but 
naturalists  have  long  felt  a  profound  conviction  that  there  is  a 
natural  system.  This  system,  it  is  now  generally  admitted,  must 
be,  as  far  as  possible,  genealogical  in  arrangement, — that  is  the 
co-descendants  of  the  same  form  must  be  kept  together  in  one 
group,  apart  from  the  co-descendants  of  any  other  form;  but  if 
the  parent-forms  are  related,  so  will  be  their  descendants,  and 
the  two  groups  together  will  form  a  larger  group.  The  amount  of 
difference  between  the  several  groups — that  is  the  amount  of 
modification  which  each  has  undergone — is  expressed  by  such 
terms  as  genera,  families,  orders,  and  classes.  As  we  have  no 
record  of  the  lines  of  descent,  the  pedigree  can  be  discovered  only 
by  observing  the  degrees  of  resemblance  between  the  beings  which 
are  to  be  classed.  For  this  object  numerous  points  of  resem- 
blance are  of  much  more  importance  than  the  amount  of  sim- 
ilarity or  dissimilarity  in  a  few  points.  If  two  languages  were 
found  to  resemble  each  other  in  a  multitude  of  words  and  points 
of  construction,  they  would  be  universally  recognized  as  having 
sprung  from  a  common  source,  notwithstanding  that  they  differed 
greatly  in  some  few  words  or  points  of  construction.  But  with 
organic  beings  the  points  of  resemblance  must  not  consist  of 
adaptations  to  similar  habits  of  life:  two  animals  may,  for  in- 
stance, have  had  their  whole  frames  modified  for  living  in  the 
water,  and  yet  they  will  not  be  brought  any  nearer  to  each  other 
in  the  natural  system.  Hence  we  can  see  how  it  is  that  resem- 
blances in  several  unimportant  structures,  in  useless  and  rudi- 
mentary organs,  or  not  now  functionally  active,  or  in  an  embry- 


3  Westwood,  'Modern  Class  of  Insects,'  vol.  ii.  1840,  p.  87. 

i 


AFFINITIES    AND    GENEALOGY.  145 

ological  condition,  are  by  far  the  most  serviceable  for  classifica- 
tion; for  they  can  hardly  be  due  to  adaptations  within  a  late 
period;  and  thus  they  reveal  the  old  lines  of  descent  or  of  true 
affinity. 

We  can  further  see  why  a  great  amount  of  modification  in 
some  one  character  ought  not  to  lead  us  to  separate  widely  any 
two  organisms.  A  part  which  already  differs  much  from  the 
same  part  in  other  allied  forms  has  already,  according  to  the 
theory  of  evolution,  varied  much;  consequently  it  would  (as  long 
as  the  organism  remained  exposed  to  the  same  exciting  condi- 
tions) be  liable  to  further  variations  of  the  same  kind;  and 
these,  if  beneficial,  would  be  preserved,  and  thus  be  continually 
augmented.  In  many  cases  the  continued  development  of  a  part, 
for  instance,  of  the  beak  of  a  bird,  or  of  the  teeth  of  a  mammal, 
would  not  aid  the  species  in  gaining  its  food,  or  for  any  other 
object;  but  with  man  we  can  see  no  definite  limit  to  the  con- 
tinued development  of  the  brain  and  mental  faculties,  as  far  as 
advantage  is  concerned.  Therefore  in  determining  the  position 
of  man  in  the  natural  or  genealogical  system,  the  extreme  de- 
velopment of  his  brain  ought  not  to  outweigh  a  multitude  of 
resemblances  in  other  less  important  or  quite  unimportant  points. 

The  greater  number  of  naturalists  who  have  taken  into  con- 
sideration the  whole  structure  of  man,  including  his  mental 
faculties,  have  followed  Blumenbach  and  Cuvier,  and  have  placed 
man  in  a  separate  Order,  under  the  title  of  the  Bimana,  and 
therefore  on  an  equality  with  the  orders  of  the  Quadrumana, 
Carnivora,  &c.  Recently  many  of  our  best  naturalists  have 
recurred  to  the  view  first  propounded  by  Linnasus,  so  remarkable 
for  his  sagacity,  and  have  placed  man  in  the  same  Order  with 
the  Quadrumana,  under  the  title  of  the  Primates.  The  justice  of 
this  conclusion  will  be  admitted:  for  in  the  first  place,  we  must 
bear  in  mind  the  comparative  insignificance  for  classification 
of  the  great  development  of  the  brain  in  man,  and  that  the 
strongly-marked  differences  between  the  skulls  of  man  and  the 
Quadrumana  (lately  insisted  upon  Bischoff,  Aeby,  and  others, 
apparently  follow  from  their  differently  developed  brains.  In 
the  second  place,  we  must  remember  that  nearly  all  the  other 
and  more  important  differences  between  man  and  the  Quadrumana 
are  manifestly  adaptive  in  their  nature,  and  relate  chiefly  to  the 
erect  position  of  man;  such  as  the  structure  of  his  hand,  foot, 
and  pelvis,  the  curvature  of  his  spine,  and  the  position  of  his 
head.  The  family  of  Seals  offers  a  good  illustration  of  the  small 
importance  of  adaptive  characters  for  classification.  These 
animals  differ  from  all  other  Carnivora  in  the  form  of  their 
bodies  and  in  the  structure  of  their  limbs,  far  more  than  does 
man  from  the  higher  apes;   yet  in  most  systems,  from  that  of 

11 


146  THE   DESCENT  OF  MAN. 

Cuvier  to  the  most  recent  one  by  Mr.  Flower/  seals  are  ranked 
as  a  mere  family  in  the  Order  of  the  Carnivora.  If  man  had  not 
been  his  own  classifier,  he  would  never  have  thought  of  founding 
a  separate  order  for  his  own  reception. 

It  would  be  beyond  my  limits,  and  quite  beyond  my  knowledge, 
even  to  name  the  innumerable  points  of  structure  in  which  man 
agrees  with  the  other  Primates.  Our  great  anatomist  and  phil- 
osopher. Prof.  Huxley,  has  fully  discussed  this  subject,^  and  con- 
cludes that  man  in  all  parts  of  his  organization  differs  less  from 
the  higher  apes,  than  these  do  from  the  lower  members  of  the 
same  group.  Consequently  there  "is  no  justification  for  placing 
"man  in  a  distinct  order." 

In  an  early  part  of  this  work  I  brought  forward  various  facts, 
showing  how  closely  man  agrees  in  constitution  with  the  higher 
mammals;  and  this  agreement  must  depend  on  our  close  sim- 
ilarity in  minute  structure  and  chemical  composition.  I  gave, 
as  instances,  our  liability  to  the  same  diseases,  and  to  the  attacks 
of  allied  parasites;  our  tastes  in  common  for  the  same  stimu- 
lants, and  the  similar  effects  produced  by  them,  as  well  as  by 
various  drugs,  and  other  such  facts. 

As  small  unimportant  points  of  resemblance  between  man  and 
the  Quadrumana  are  not  commonly  noticed  in  systematic  works, 
and  as,  when  numerous,  they  clearly  reveal  our  relationship,  I 
will  specify  a  few  such  points.  The  relative  position  of  our  fea- 
tures is  manifestly  the  same;  and  the  various  emotions  are  dis- 
played by  nearly  similar  movements  of  the  muscles  and  skin, 
chiefly  above  the  eyebrows  and  round  the  mouth.  Some  few 
expressions  are,  indeed,  almost  the  same,  as  in  the  weeping  of 
certain  kinds  of  monkeys  and  in  the  laughing  noise  made  by 
others,  during  which  the  corners  of  the  mouth  are  drawn  back- 
wards, and  the  lower  eyelids  wrinkled.  The  external  ears  are 
curiously  alike.  In  man  the  nose  is  much  more  prominent  than 
in  most  monkeys;  but  we  may  trace  the  commencement  of  an 
aquiline  curvature  in  the  nose  of  the  Hoolock  Gibbon;  and  this 
in  the  Semnopithecus  nasica  is  carried  to  a  ridiculous  extreme. 

The  faces  of  many  monkeys  are  ornamented  with  beards,  whis- 
kers, or  moustaches.  The  hair  on  the  head  grows  to  a  great 
length  in  some  species  of  Semnopithecus;^  and  in  the  Bonnet 
monkey  (Macacus  radiatus)  it  radiates  from  a  point  on  the  crown, 
with  a  parting  down  the  middle.  It  is  commonly  said  that  the 
forehead  gives  to  man  his  noble  and  intellectual  appearance;  but 
the  thick  hair  on  the  head  of  the  Bonnet  monkey  terminates 
downwards  abruptly,  and  is  succeeded  by  hair  so  short  and  fine 

*  'Proc.  Zoolog.  Soc'  1S63,  p.  4. 

s  'Evidence  as  to  Man's  Place  in  Nature,'  1863,  p.  70,  et  passim. 

« Isid.  Geoffrey,  'Hist.  Nat.  Gen.'  torn.  ii.  1859,  p.  217. 


Ai^FINITIES    AND    GENEALOGY.  147 

that  at  a  little  distance  the  forehead,  with  the  exception  of  the 
eyebrows,  appears  quite  naked.  It  has  been  erroneously  asserted 
that  eyebrows  are  not  present  in  any  monkey.  In  the  species 
just  named  the  degree  of  nakedness  of  the  forehead  differs  in 
different  individuals;  and  Eschricht  states'  that  in  our  children 
the  limit  between  the  hairy  scalp  and  the  naked  forehead  is 
sometimes  not  well  defined;  so  that  here  we  seem  to  have  a 
trifling  case  of  reversion  to  a  progenitor,  in  whom  the  forehead 
had  not  as  yet  become  quite  naked. 

It  is  well  known  that  the  hair  on  our  arms  tends  to  converge 
from  above  and  below  to  a  point  at  the  elbow.  This  curious 
arrangement,  so  unlike  that  in  most  of  the  lower  mammals,  is 
common  to  the  gorilla,  chimpanzee,  orang,  some  species  of  Hy- 
lobates,  and  even  to  some  few  American  monkeys.  But  in  Hylo- 
bates  agilis  the  hair  on  the  fore-arm  is  directed  downwards  or 
towards  the  wrist  in  the  ordinary  manner;  and  in  H.  lar  it  is 
nearly  erect,  with  only  a  very  slight  forward  inclination;  so  that 
in  this  latter  species  it  is  in  a  transitional  state.  It  can  hardly 
be  doubted  that  with  most  mammals  the  thickness  of  the  hair  on 
the  back  and  its  direction,  is  adapted  to  throw  off  the  rain;  even 
the  transverse  hairs  on  the  fore-legs  of  a  dog  may  serve  for  this 
end  when  he  is  coiled  up  asleep.  Mr.  Wallace,  who  has  carefully 
studied  the  habits  of  the  orang,  remarks  that  the  convergence  of 
the  hair  towards  the  elbow  on  the  arms  of  the  orang  may  be 
explained  as  serving  to  throw  off  the  rain,  for  this  animal  during 
rainy  weather  sits  with  its  arms  bent,  and  with  the  hands  clasped 
round  a  branch  or  over  its  head.  According  to  Livingstone,  the 
gorilla  also  "sits  in  pelting  rain  with  his  hands  over  his  head."^.. 
If  the  above  explanation  is  correct,  as  seems  probable,  the  direc-  , 
tion  of  the  hair  on  our  own  arms  offers  a  curious  record  of  our/ 
former  state;  for  no  one  supposes  that  it  is  now  of  any  use  in 
throwing  off  the  rain;  nor,  in  our  present  erect  condition,  is  it 
properly  directed  for  this  purpose. 

It  would,  however,  be  rash  to  trust  too  much  to  the  principle 
of  adaptation  in  regard  to  the  direction  of  the  hair  in  man  or  his 
early  progenitors;  for  it  is  itnpossible  to  study  the  figures  given 
by  Eschricht  of  the  arrangement  of  the  hair  on  the  human  foetus 
(this  being  the  same  as  in  the  adult)  and  not  agree  with  this 
excellent  observer  that  other  and  more  complex  causes  have 
intervened.  The  points  of  convergence  seem  to  stand  in  some 
relation  to  those  points  in  the  embryo  which  are  last  closed  in 
during  development.    There  appears,  also,  to  exist  some  relation 

'  'Ueber  die  Richtung  der  Haare,'  &c.,  Muller's  'Archiv  fur  Anat.  und 
Phys'  1837,  s.   51. 
8  Quoted  by  Reade,  'The  African  Sketch  Book,'  vol.  1.  1873,  p.  152. 


148  THE  DESCENT  OF  MAN. 

between  the  arrangement  of  the  hair  on  the  limbs,  and  the  course 
of  the  medullary  arteries.'' 

It  must  not  be  supposed  that  the  resemblances  between  man 
and  certain  apes  in  the  above  and  many  other  points — such  as  in 
having  a  naked  forehead,  long  tresses  on  the  head,  «S:c. — are  all 
necessarily  the  result  of  unbroken  inheritance  from  a  common 
progenitor,  or  of  subsequent  reversion.  Many  of  these  resem- 
blances are  more  probably  due  to  analogous  variation,  which 
follows,  as  I  have  elsewhere  attempted  to  show,^"  from  co-de- 
scended organisms  having  a  similar  constitution,  and  having  been 
acted  on  by  like  causes  inducing  similar  modifications.  With  re- 
spect to  the  similar  direction  of  the  hair  on  the  fore-arms  of  man 
and  certain  monkeys,  as  this  character  is  common  to  almost  all 
the  anthropomorphous  apes,  it  may  probably  be  attributed  to  in- 
heritance; but  this  is  not  certain,  as  some  very  distinct  American 
monkeys  are  thus  characterized. 

Although,  as  we  have  now  seen,  man  has  no  jiist  right  to  form 
a  separate  Order  for  his  own  reception,  he  may  perhaps  claim  a 
distinct  Sub-order  or  Family.  Prof.  Huxley,  in  his  last  work," 
divides  the  Primates  into  three  Sub-orders;  namely,  the  An- 
thropidse  with  man  alone,  the  Simiadae  including  monkeys  of  all 
kinds,  and  the  Lemuridas  with  the  diversified  genera  of  lemurs. 
As  far  as  differences  in  certain  important  points  of  structure  are 
concerned,  man  may  no  doubt  rightly  claim  the  rank  of  a  Sub- 
order; and  this  rank  is  too  low  if  we  look  chiefiy  to  his  mental 
faculties.  Nevertheless,  from  a  genealogical  point  of  view  it 
appears  that  this  rank  is  too  high,  and  that  man  ought  to  form 
merely  a  Family,  or  possibly  even  only  a  Sub-family.  If  we 
imagine  three  lines  of  descent  proceeding  from  a  common  stock, 
it  is  quite  conceivable  that  two  of  them  might  after  the  lapse  of 
ages  be  so  slightly  changed  as  still  to  remain  as  species  of  the 
same  genus,  whilst  the  third  line  might  become  so  greatly  modi- 
fied as  to  deserve  to  rank  as  a  distinct  Sub-family,  Family,  or 
even  Order.  But  in  this  case  it  is  almost  certain  that  the  third 
line  would  still  retain  through  inheritance  numerous  small  points 
of  resemblance  with  the  other  two.  Here,  then,  would  occur 
the  difficulty,  at  present  insoluble,  how  much  weight  we  ought 
to  assign  in  our  classifications  to  strongly-marked  differences  in 
some  few  points, — that  is,  to  the  amount  of  modification  under- 

»  On  the  hair  in  Hylobates,  see  'Nat.  Hist,  of  Mammals,'  by  C.  L. 
Martin,  1841,  p.  415.  Also,  Isid.  Geoffroy  on  the  American  monkeys  and 
other  kinds,  'Hist.  Nat.  Gen.'  vol.  ii.  1859,  p.  216,  243.  Eschricht  ibid. 
s,  46,  55,  61.  Owen,  'Anat.  of  Vertebrates,'  vol.  iii.  p.  619.  Wallace,  'Con- 
tributions to  the  Theory  of  Natural  Selection,'  1870,  p.  344. 

10  'Origin  of  Species,'  5th  edit.  1S69,  p.  194.  'The  Variation  of  Animals 
and  Plants  under  Domestication,'  vol.  ii.  1868,  p.  348. 

u  'An  Introduction  to  the  Classification  of  Animals,'  1869,  p.  99. 


AFFINITIES    AND    GENEALOGY.  149 

gone;  and  how  much  to  close  resemblance  in  numerous  unim- 
portant points,  as  indicating  the  lines  of  descent  or  genealogy. 
To  attach  much  weight  to  the  few  but  strong  differences  is  the 
most  obvious  and  perhaps  the  safest  course,  though  it  appears 
more  correct  to  pay  great  attention  to  the  many  small  resem- 
blances, as  giving  a  truly  natural  classification. 

In  forming  a  judgment  on  this  head  with  reference  to  man,  we 
must  glance  at  the  classification  of  the  Simiadse.  This  family  is 
divided  by  almost  all  naturalists  into  the  Catarhine  group,  or 
Old  World  monkeys,  all  of  which  are  characterized  (as  their 
name  expresses)  by  the  peculiar  structure  of  their  nostrils,  and  by 
having  four  premolars  in  each  jaw;  and  into  the  Platyrhine 
group  or  New  World  monkeys  (including  two  very  distinct  sub- 
groups), all  of  which  are  characterized  by  differently  constructed 
nostrils,  and  by  having  six  premolars  in  each  jaw.  Some  other 
small  differences  might  be  mentioned.  Now  man  unquestionably 
belongs  in  his  dentition,  in  the  structure  of  his  nostrils,  and 
some  other  respects,  to  the  Catarhine  or  Old  World  division;  nor 
does  he  resemble  the  Platyrhines  more  closely  than  the  Catar- 
hines  in  any  characters,  excepting  in  a  few  of  not  much  import- 
ance and  apparently  of  an  adaptive  nature.  It  is  therefore  against 
all  probability  that  some  New  World  species  should  have  for- 
merly varied  and  produced  a  man-like  creature,  with  all  the  dis- 
tinctive characters  proper  to  the  Old  World  division;  losing  at 
the  same  time  all  its  own  distinctive  characters.  There  can,  con- 
sequently, hardly  be  a  doubt  that  man  is  an  off-shoot  from  the 
Old  World  Simian  stem;  and  that  under  a  genealogical  point  of 
view,  he  must  be  classed  with  the  Catarhine  division.^^ 

The  anthropomorphous  apes,  namely,  the  gorilla,  chimpanzee, 
orang,  and  hylobates,  are  by  most  naturalists  separated  from  the 
other  Old  World  monkeys,  as  a  distinct  sub-group.  I  am  aware 
that  Gratiolet,  relying  on  the  structure  of  the  brain,  does  not 
admit  the  existence  of  this  sub-group,  and  no  doubt  it  is  a  broken 
one.  Thus  the  orang,  as  Mr.  St.  G.  Mivart  remarks,^^  "is  one  of 
"the  most  peculiar  and  aberrant  forms  to  be  found  in  the  Order." 
The  remaining  non-anthropomorphous  Old  World  monkeys,  are 
again  divided  by  some  naturalists  into  two  or  three  smaller  sub- 
groups; the  genus  Semnopithecus,  -mth  its  peculiar  sacculated 
stomach,  being  the  type  of  one  such  sub-group.     But  it  appears 

12  This  is  nearly  the  same  classification  as  that  provisionally  adopted 
by  Mr.  St.  George  Mivart  ('Transact.  Pliilosoph.  Soc'  1867,  p.  300),  who, 
after  separating  the  Lemuridae,  divides  the  remainder  of  the  Primates 
into  the  Hominidae,  the  Simiadae  which  answer  to  the  Catarhines,  the 
Cebidae,  and  the  Hapalidae,— these  two  latter  groups  ansv/ering-  to  the 
Platyrhines.  Mr.  Mivart  still  abides  by  the  same  view;  see  'Nature,' 
1871,  p.  4S1. 

13  'Transact.  Zoolog.  Soc'  vol.  vi.  1867,  p.  214, 

11 


150  THE  DESCENT  OF  MAN. 

from  M.  Gaudry's  wonderful  discoveries  in  Attica,  that  during 
the  Miocene  period  a  form  existed  there,  which  connected  Sem- 
nopithecus  and  Macacus;  and  this  probably  illustrates  the  man- 
ner in  which  the  other  and  higher  groups  were  once  blended  to- 
gether. 

If  the  anthropomorphous  apes  be  admitted  to  form  a  natural 
sub-group,  then  as  man  agrees  with  them,  not  only  in  all  those 
characters  which  he  possesses  in  common  with  the  whole  Catar- 
hine  group,  but  in  other  peculiar  characters,  such  as  the  absence 
of  a  tail  and  of  callosities,  and  in  general  appearance,  we  may 
infer  that  some  ancient  member  of  the  anthropomorphous  sub- 
group gave  birth  to  man.  It  is  not  probable  that,  through  the 
law  of  analogous  variation,  a  member  of  one  of  the  other  lower 
sub-groups  should  have  given  rise  to  a  man-like  creature,  re- 
sembling the  higher  anthropomorphous  apes  in  so  many  respects. 
No  doubt  man,  in  comparison  with  most  of  his  allies,  has  under- 
gone an  extraordinary  amount  of  modification,  chiefly  in  conse- 
quence of  the  great  development  of  his  brain  and  his  erect  po- 
sition; nevertheless,  we  should  bear  in  mind  that  he  "is  but  one 
"of  several  exceptional  forms  of  Primates."^* 

Every  naturalist,  who  believes  in  the  principle  of  evolution, 
will  grant  that  the  two  main  divisions  of  the  Simiadae,  namely 
the  Catarhine  and  Platyrhine  monkeys,  with  their  sub-groups, 
have  all  proceeded  from  some  one  extremely  ancient  progenitor, 
The  early  descendants  of  this  progenitor,  before  they  had  diverged 
to  any  considerable  extent  from  each  other,  would  still  have 
formed  a  single  natural  group;  but  some  of  the  species  or  incip- 
ient genera  would  have  already  begun  to  indicate  by  their  diverg- 
ing characters  the  future  distinctive  marks  of  the  Catarhine  and 
Platyrhine  divisions.  Hence  the  members  of  this  supposed  an- 
cient group  would  not  have  been  so  uniform  in  their  dentition, 
or  in  the  structure  of  their  nostrils,  as  are  the  existing  Catarhine 
monkeys  in  one  way  and  the  Platyrhines  in  another  way,  but 
would  have  resembled  in  this  respect  the  allied  Lemuridse,  which 
differ  greatly  from  each  other  in  the  form  of  their  muzzles,^^ 
and  to  an  extraordinary  degree  in  their  dentition. 

The  Catarhine  and  Platyrhine  monkeys  agree  in  a  multitude  of 
characters,  as  is  shown  by  their  unquestionably  belonging  to  one 
and  the  same  Order.  The  many  characters  which  they  possess 
in  common  can  hardly  have  been  independently  acquired  by  so 
many  distinct  species;  so  that  these  characters  must  have  been 
inherited.  But  a  naturalist  would  undoubtedly  have  ranked  as 
an  ape  or  a  monkey,  an  ancient  form  which  possessed  many  char- 

"Mr.  St.  G.  Mivart,  'Transact.  Phil.  Soc'  1867,  p.  410. 
"  Messrs.   Murie  and  Mivart  on  the  Lemuroidea,   'Transact.  Zoologr. 
Soc.'  vol.  vii.  1869,  p.  5. 


AFFINITIES    AND    GENEALOGY.  151 

acters  common  to  the  Catarhine  and  Platyrhine  monkeys,  other 
characters  in  an  intermediate  condition,  and  some  few,  perhaps, 
distinct  from  those  now  found  in  either  group.  And  as  man 
from  a  genealogical  point  of  view  belongs  to  the  Catarhine  or 
Old  World  stock,  we  must  conclude,  however  much  the  conclusion 
may  revolt  our  pride,  that  our  early  progenitors  would  have 
been  properly  thus  designated.^^  But  we  must  not  fall  into  the 
error  of  supposing  that  the  early  progenitor  of  the  whole  Simian 
stock,  including  man,  was  identical  with,  or  even  closely  re- 
sembled, any  existing  ape  or  monkey. 

On  the  Birthplace  and  Antiquity  of  Man.— We  are  naturally 
led  to  enquire,  where  was  the  birthplace  of  man  at  that  stage  of 
descent  when  our  progenitors  diverged  from  the  Catarhine  stock? 
The  fact  that  they  belonged  to  this  stock  clearly  shows  that 
they  inhabited  the  Old  World;  but  not  Australia  nor  any  oceanic 
island,  as  we  may  infer  from  the  laws  of  geographical  distribu- 
tion. In  each  great  region  of  the  world  the  living  mammals  are 
closely  related  to  the  extinct  species  of  the  same  region.  It  is 
therefore  probable  that  Africa  was  formerly  inhabited  by  extinct 
apes  closely  allied  to  the  gorilla  and  chimpanzee;  and  as  these  two 
species  are  now  man's  nearest  allies,  it  is  somewhat  more  probable 
that  our  early  progenitors  lived  on  the  African  continent  than 
elsewhere.  But  it  is  useless  to  speculate  on  this  subject;  for  two 
or  three  anthropomorphous  apes,  one  the  Dryopithecus"  of  Lartet, 
nearly  as  large  as  a  man,  and  closely  allied  to  Hylobates,  existed 
in  Europe  during  the  Miocene  age;  and  since  so  remote  a  period 
the  earth  has  certainly  undergone  many  great  revolutions,  and 
there  has  been  ample  time  for  migration  on  the  largest  scale. 

At  the  period  and  place,  whenever  and  wherever  it  was,  when 
man  first  lost  his  hairy  covering,  he  probably  inhabited  a  hot 
country;  a  circumstance  favorable  for  the  frugiferous  diet  on 
which,  judging  from  analogy,  he  subsisted.  We  are  far  from 
knowing  how  long  ago  it  was  when  man  first  diverged  from  the 
Catarhine  stock;  but  it  may  have  occurred  at  an  epoch  as  remote 
as  the  Eocene  period;  for  that  the  higher  apes  had  diverged 
from  the  lower  apes  as  early  as  the  Upper  Miocene  period  is 
shown  by  the  existence  of  the  Dryopithecus.  We  are  also  quite 
ignorant  at  how  rapid  a  rate  organisms,  v/hether  high  or  low 
in  the  scale,  may  be  modified  under  favorable  circumstances;  we 

1^  Hackel  has  come  to  this  same  conclusion.  See  'Ueber  die  Entste- 
htmg-  des  Menschengeschlechts,'  in  Virchow's  'Sammlung'.  g-emein. 
wissen.  Vortrage,'  1868,  s.  61.  Also  his  'Naturliche  Schopfungsg-e- 
schichte,'  1868,  in  which  he  gives  in  detail  his  views  on  the  genealogy 
of  man. 

17  Dr.  C.  Forsyth  Major,  'Sur  les  Singes  Fossiles  trouves  on  Italia;' 
'Soc.  Ital.  des  So.  Nat.'  torn.  xv.  1872. 


152  THE  DESCENT  OF  MAN. 

knoAv,  however,  that  some  have  retained  the  same  form  during 
an  enormous  lapse  of  time.  From  what  we  see  going  on  under 
domestication,  we  learn  that  some  of  the  co-descendants  of  the 
same  species  may  he  not  at  all,  some  a  little,  and  some  greatly 
changed,  all  within  the  same  period.  Thus  it  may  have  been 
with  man,  who  has  undergone  a  great  amount  of  modification  in 
certain  characters  in  comparison  with  the  higher  apes. 

The  great  break  in  the  organic  chain  between  man  and  his 
nearest  allies,  which  cannot  be  bridged  over  by  any  extinct  or 
living  species,  has  often  been  advanced  as  a  grave  objection  to 
the  belief  that  man  is  descended  from  some  lower  form;  but  this 
objection  will  not  appear  of  much  weight  to  those  who,  from 
general  reasons,  believe  in  the  general  principle  of  evolution. 
Breaks  often  occur  in  all  parts  of  the  series,  some  being  wide, 
sharp  and  defined,  others  less  so  in  various  degrees;  as  between 
the  orang  and  its  nearest  allies — between  the  Tarsius  and  the 
other  Lemuridse — between  the  elephant,  and  in  a  more  striking 
manner  between  the  Ornithorhynchus  or  Echidna,  and  all  other 
mammals.  But  these  breaks  depend  merely  on  the  number  of 
related  forms  which  have  become  extinct.  At  some  future  period, 
not  very  distant  as  measured  by  centuries,  the  civilized  races 
of  man  will  almost  certainly  exterminate,  and  replace,  the  savage 
races  throughout  the  world.  At  the  same  time  the  anthropomor- 
phous apes,  as  Professor  Schaaffhausen  has  remarked,^^  will  no 
doubt  be  exterminated.  The  break  between  man  and  his  nearest 
allies  will  then  be  wider,  for  it  will  intervene  between  man  in 
a  more  civilized  state,  as  we  may  hope,  even  than  the  Caucasian, 
and  some  ape  as  low  as  a  baboon,  instead  of  as  now  between  the 
negro  or  Australian  and  the  gorilla. 

With  respect  to  the  absence  of  fossil  remains,  serving  to  con- 
nect man  with  his  ape-like  progenitors,  no  one  will  lay  much 
stress  on  this  fact  who  reads  Sir  C.  Lyell's  discussion,^^  where 
he  shows  that  in  all  the  vertebrate  classes  the  discovery  of  fossil 
remains  has  been  a  very  slow  and  fortuitous  process.  Nor 
should  it  be  forgotten  that  those  regions  which  are  the  most 
likely  to  afford  remains  connecting  man  with  some  extinct  ape- 
like creature,  have  not  as  yet  been  searched  by  geologists. 

Lower  Stages  in  the  Genealogy  of  Man. — We  have  seen  that 
man  appears  to  have  diverged  from  the  Catarhines  or  Old  World 
division  of  the  Simiadse,  after  these  had  diverged  from  the  New 
World  division.  We  will  now  endeavor  to  follow  the  remote 
traces  of  his  genealogy,  trusting  principally  to  the  mutual  affin- 


18  'Anthropolog-ical  Review,'  April,  1SG7,  p.  236. 

19  'Elements  of  Geology,'    1865,   pp.   5S3-585.    'Antiquity   of  Man,'   1863, 
p.  145. 


JLFTINITIES   AND   GENEALOGY.  153 

ities  between  the  various  classes  and  orders,  witli  some  slight 
reference  to  the  periods,  as  far  as  ascertained,  of  their  suc- 
ce.-!sive  appearance  on  the  earth.  The  Lemuridas  stand  below 
and  near  to  the  Simiadee,  and  constitute  a  very  distinct  family 
of  the  Primates,  or,  according  to  Hackel  and  others,  a  distinct 
Order.  This  group  is  diversified  and  broken  to  an  extraordinary 
degree,  and  includes  many  aberrant  forms.  It  has,  therefore, 
probably  suffered  much  extinction.  Most  of  the  remnants  sur- 
vive on  islands,  such  as  Madagascar  and  the  Malayan  archipelago, 
where  they  have  not  been  exposed  to  so  severe  a  competition  as 
they  would  have  been  on  well-stocked  continents.  This  group 
likewise  presents  many  gradations,  leading,  as  Huxley  remarks,^^ 
"insensibly  from  the  crown  and  summit  of  the  animal  creation 
"down  to  creatures  from  which  there  is  but  a  step,  as  it  seems, 
"to  the  lowest,  smallest,  and  least  intelligent  of  the  placental 
"mammalia."  From  these  various  considerations  it  is  probable 
that  the  Simiadae  were  originally  developed  from  the  progenitors 
of  the  existing  Lemuridas;  and  these  in  their  turn  from  forms 
standing  very  low  in  the  mammalian  series. 

The  Marsupials  stand  in  many  important  characters  below  the 
placental  mammals.  They  appeared  at  an  earlier  geological 
period,  and  their  range  was  formerly  much  more  extensive  than 
at  present.  Hence  the  Placentata  are  generally  supposed  to  have 
been  derived  from  the  Implacentata  or  Marsupials;  not,  however, 
from  forms  closely  resembling  the  existing  Marsupials,  but  from 
their  early  progenitors.  The  Monotremata  are  plainly  allied  to 
the  Marsupials,  forming  a  third  and  still  lower  division  in  the 
great  mammalian  series.  They  are  represented  at  the  present 
day  solely  by  the  Ornithorhynchus  and  Echidna;  and  these  two 
forms  may  be  safely  considered  as  relics  of  a  much  larger  group 
representatives  of  which  have  been  preserved  in  Australia  through 
some  favorable  concurrence  of  circumstances.  The  Monotremata 
are  eminently  interesting,  as  leading  in  several  important  points 
of  structure  towards  the  class  of  reptiles. 

In  attempting  to  trace  the  genealogy  of  the  Mammalia,  and 
therefore  of  man,  lower  down  in  the  series,  we  become  involved 
in  greater  and  greater  obscurity;  but  as  a  most  capable  judge, 
Mr.  Parker,  has  remarked,  we  have  good  reason  to  believe,  that 
no  true  bird  or  reptile  intervenes  in  the  direct  line  of  descent. 
He  who  wishes  to  see  what  ingenuity  and  knowledge  can  effect, 
may  consult  Prof.  Hackel's  works.^^    I  will  content  myself  with 

20  'Man's  Place  in  Nature,'  p.  105. 

-1  Elaborate  tables  are  given  in  his  'Generelle  Morphologie'  (B.  li. 
s.  cliii.  and  s.  425);  and  with  more  especial  reference  to  man  in  his 
'Naturliche  Schopfungschichte,'  1868.  Prof.  Huxley,  in  reviewing 
this  latter  work  i('The  Academy,'  1869,  p.  42)  says,  that  he  considers  the 
phylum  or  lines  of  descent  of  the  Vertebrata  to  be  admirably  discussed 


154  THE   DESCENT  OF  MAN. 

a  few  general  remarks.  Every  evolutionist  will  admit  that  the 
five  great  vertebrate  classes,  namely,  mammals,  birds,  reptiles, 
amphibians,  and  fishes,  are  descended  from  some  one  prototype; 
for  they  have  much  in  common,  especially  during  their  embryonic 
state.  As  the  class  of  fishes  is  the  most  lowly  organized,  and 
appeared  before  the  others,  we  may  conclude  that  all  the  members 
of  the  vertebrate  kingdom  are  derived  from  some  fish-like  animal. 
The  belief  that  animals  so  distinct  as  a  monkey,  an  elephant,  a 
humming-bird,  a  snake,  a  frog,  and  a  fish,  &c.,  could  all  have 
sprung  from  the  same  parents,  will  appear  monstrous  to  those 
who  have  not  attended  to  the  recent  progress  of  natural  history. 
For  this  belief  Implies  the  former  existence  of  .links  binding 
closely  together  all  these  forms,  now  so  utterly  unlike. 

Nevertheless,  it  is  certain  that  groups  of  animals  have  existed, 
or  do  now  exist,  which  serve  to  connect  several  of  the  great 
vertebrate  classes  more  or  less  closely.  We  have  seen  that  the 
Ornithorhynchus  graduates  towards  reptiles;  and  Prof.  Huxley 
has  discovered,  and  is  confirmed  by  Mr.  Cope  and  others,  that 
the  Dinosaurians  are  in  many  important  characters  intermediate 
between  certain  reptiles  and  certain  birds — the  birds  referred 
to  being  the  ostrich-tribe  (itself  evidently  a  widely-diffused 
remnant  of  a  larger  group)  and  the  Archeopteryx,  that  strange 
Secondary  bird,  with  a  long  lizard-like  tail.  Again,  according  to 
Prof.  Owen,"  the  Ichthyosaurians — great  sea-lizards  furnished 
with  paddles — present  many  affinities  with  fishes,  or  rather 
according  to  Huxley,  with  amphibians;  a  class  which,  including 
in  its  highest  division  frogs  and  toads,  is  plainly  allied  to  the 
Ganoid  fishes.  These  latter  fishes  swarmed  during  the  earlier 
geological  periods,  and  were  constructed  on  what  is  called  a 
generalized  type,  that  is,  they  presented  diversified  affinities  with 
other  groups  of  organisms.  The  Lepidosiren  is  also  so  closely 
allied  to  amphibians  and  fishes,  that  naturalists  long  disputed  in 
which  of  these  two  classes  to  rank  it;  it,  and  also  some  few 
Ganoid  fishes,  have  been  preserved  from  utter  extinction  by 
inhabiting  rivers,  which  are  harbors  of  refuge,  and  are  related 
to  the  great  waters  of  the  ocean  in  the  same  way  that  islands 
are  to  continents. 

Lastly,  one  single  member  of  the  immense  and  diversified  class 
of  fishes,  namely,  the  lancelet  or  amphioxus,  is  so  different  from 
all  other  fishes,  that  Hackel  maintains  that  it  ought  to  form  a 
distinct  class  in  the  vertebrate  kingdom.  This  fish  is  remarkable 
for  its  negative  characters;  it  can  hardly  be  said  to  possess  a 
brain,  vertebral  column,  or  heart,  &c.;    so  that  it  was  classed  by 

by  Hackel,   althoug-h   he   differs  on  some  points.    He  expresses,    also, 
his  high  estimate  of  the  general  tenor  and  spirit  of  the  whole  work. 
22  'Palaeontology,'  1860,  p.  199. 


AFFINITIES   AND    GENEALOGY.  155 

the  older  naturalists  amongst  the  worms.  Many  years  ago  Prof. 
Goodsir  perceived  that  the  lancelet  presented  some  affinities  with 
the  Ascidians,  which  are  invertebrate,  hermaphrodite,  marine 
creatures  permanently  attached  to  a  support.  They  hardly  ap- 
pear like  animals,  and  consist  of  a  simple,  tough,  leathery  sack, 
with  two  small  projecting  orifices.  They  belong  to  the  Mollus- 
coida  of  Huxley— a  lower  division  of  the  great  kingdom  of  the 
Mollusca;  but  they  have  recently  been  placed  by  some  natural- 
ists amongst  the  Vermes  or  worms.  Their  larvae  somewhat 
resemble  tadpoles  in  shape,-^  and  have  the  power  of  swimming 
freely  about.  M.  Kovalevsky-*  has  lately  observed  that  the  larvae 
of  Ascidians  are  related  to  the  Vertebrata,  in  their  manner  of 
development,  in  the  relative  position  of  the  nervous  system, 
and  in  possessing  a  structure  closely  like  the  chorda  dorsalis  of 
vertebrate  animals;  and  in  this  he  has  been  since  confirmed  by 
Prof.  Kupffer.  M.  Kovalevsky  writes  to  me  from  Naples,  that 
he  has  now  carried  these  observations  yet  further;  and  should 
his  results  be  well  established,  the  whole  will  form  a  discovery 
of  the  very  greatest  value.  Thus,  if  we  may  rely  on  embryology, 
ever  the  safest  guide  in  classification,  it  seems  that  we  have  at 
last  gained  a  clue  to  the  source  whence  the  Vertebrata  were  de- 
rived.^^  We  should  then  be  justified  in  believing  that  at  an  ex- 
tremely remote  period  a  group  of  animals  existed,  resembling  in 
many  respects  the  larvae  of  our  present  Ascidians,  which  diverged 
into  two  great  branches — the  one  retrograding  in  development 
and  producing  the  present  class  of  Ascidians,  the  other  rising  to 
the  crown  and  summit  of  the  animal  kingdom  by  giving  birth 
to  the  Vertebrata. 


23  At  the  Falkland  Islands  I  had  the  satisfaction  of  seeing,  in  April, 
1833,  and  therefore  some  years  before  any  other  naturalist,  the  loco- 
motive larvae  of  a  compound  Ascidian,  closely  allied  to  Synoicum, 
but  apparently  generically  distinct  from  it.  The  tail  was  about  five 
times  as  long-  as  the  oblong  head,  and  terminated  in  a  very  fine  fila- 
ment. It  was,  as  sketched  by  me  under  a  simple  microscope  plainly 
divided  by  transverse  opaque  partitions,  which  I  presume  represent 
the  great  cells  figured  by  Kovalevsky.  At  an  early  stage  of  develop- 
ment the  tail  was  closely  coiled  round  the  head  of  the  larva. 

24  'Memoires  de  I'Acad.  des  Sciences  de  St.  Petersbourg,'  tom.  x. 
No.  15,  1S66. 

25  But  I  am  bound  to  add  that  some  competent  judges  dispute  this 
conclusion;  for  instance,  M.  Giard,  in  a  series  of  papers  in  the  Arch- 
ives de  Zoologie  Experimentale,'  for  1872.  Nevertheless,  this  natural- 
ist remarks,  p.  281,  "L'organisation  de  la  larve  ascidienne  en  dehors 
"de  toute  hypothese  et  de  toute  theorie,  nous  montre  comment  la 
"nature  peut  produire  la  disposition  fondamentale  du  type  vertebre 
"(I'existence  d'une  corde  dorsale)  chez  un  invertebre  par  la  seule  con- 
"dition  vitale  de  I'ad.aptation,  et  cette  simple  possibilite  du  passage 
"supprime  I'abime  entre  les  deux  sous-regnes,  encore  bien  qu'en  ignore 
"par  ou  le  passage  s'est  fait  en  realite." 


156  THE   DESCENT  OF  MAN. 

We  have  thus  far  endeavored  rudely  to  trace  the  genealogy 
of  the  Vertebrata  by  the  aid  of  their  mutual  affinities.  We  will 
now  look  to  man  as  he  exists;  and  we  shall,  I  think,  be  able 
partially  to  restore  the  structure  of  our  early  progenitors,  during 
successive  periods,  but  not  in  due  order  of  time.  This  can  be 
effected  by  means  of  the  rudiments  which  man  still  retains,  by 
the  characters  which  occasionally  make  their  appearance  in  him 
through  reversion,  and  by  the  aid  of  the  principles  of  morphology 
and  embryology.  The  various  facts,  to  which  I  shall  here  allude, 
have  been  given  in  the  previous  chapters. 

The  early  progenitors  of  man  must  have  been  once  covered 
with  hair,  both  sexes  having  beards;  their  ears  were  probably 
pointed,  and  capable  of  movement;  and  their  bodies  were  pro- 
vided with  a  tail,  having  the  proper  muscles.  Their  limbs  and 
bodies  were  also  acted  on  by  many  muscles  which  now  only 
occasionally  reappear,  but  are  normally  present  in  the  Quadru- 
mana.  At  this  or  some  earlier  period,  the  great  artery  and  nerve 
of  the  humerus  ran  through  a  supra-condyloid  foramen.  The 
intestine  gave  forth  a  much  larger  diverticulum  or  caecum  than 
that  now  existing.  The  foot  was  then  prehensile,  judging  from 
the  condition  of  the  great  toe  in  the  foetus;  and  our  progenitors, 
no  doubt,  were  arboreal  in  their  habits,  and  frequented  some 
warm,  forest-clad  land.  The  males  had  great  canine  teeth,  which 
served  them  as  formidable  weapons.  At  a  much  earlier  period 
the  uterus  was  double;  the  excreta  were  voided  through  a  cloaca; 
and  the  eye  was  protected  by  a  third  eyelid  or  nictitating  mem- 
brane. At  a  still  earlier  period  the  progenitors  of  man  must  have 
been  aquatic  in  their  habits;  for  morphology  plainly  tells  us  that 
our  lungs  consist  of  a  modified  swim-bladder,  which  once  served 
as  a  float.  The  clefts  on  the  neck  in  the  embryo  of  man  show 
where  the  branchiae  once  existed.  In  the  lunar  or  weekly  re- 
current periods  of  some  of  our  functions  we  apparently  still  retain 
traces  of  our  primordial  birthplace,  a  shore  washed  by  the  tides. 
At  about  this  same  early  period  the  true  kidneys  were  replaced 
by  the  corpora  wolffiana.  The  heart  existed  as  a  simple  pulsating 
vessel;  and  the  chorda  dorsalis  took  the  place  of  a  vertebral 
column.  These  early  ancestors  of  man,  thus  seen  in  the  dim 
recesses  of  time,  must  have  been  as  simply,  or  even  still  more 
simply  organized  than  the  lancelet  or  amphioxus. 

There  is  one  other  point  deserving  a  fuller  notice.  It  has  long 
been  known  that  in  the  vertebrate  kingdom  one  sex  bears  rudi- 
ments of  various  accessory  parts,  appertaining  to  the  reproduc- 
tive system,  which  properly  belong  to  the  opposite  sex;  and  it 
has  now  been  ascertained  that  at  a  very  early  embryonic  period 
both  sexes  possess  true  male  and  female  glands.  Hence  some  re- 
mote progenitor  of  the  whole  vertebrate  kingdom  appears  to  have 


AFFINITIES   AND    GENEALOGY.  157 

been  hermaphrodite  or  androgynous.'^  But  here  we  encounter 
a  singular  difficulty.  In  the  mammalian  class  the  males  possess 
rudiments  of  a  uterus  with  the  adjacent  passage,  in  their  vesiculse 
prostaticse;  they  bear  also  rudiments  of  mammas,  and  some  male 
Marsupials  have  traces  of  a  marsupial  sack.-'  Other  analogous 
facts  could  be  added.  Are  we,  then,  to  suppose  that  some  ex-' 
tremely  ancient  mammal  continued  androgynous,  after  it  had 
acquired  the  chief  distinctions  of  its  class,  and  therefore  after  it 
had  diverged  from  the  lower  classes  of  the  vertebrate  kingdom? 
This  seems  very  improbable,  for  we  have  to  look  to  fishes,  the 
lowest  of  all  the  classes,  to  find  any  still  existent  androgynous 
forms.'*  That  various  accessory  parts,  proper  to  each  sex,  are 
found  in  a  rudimentary  condition  in  the  opposite  sex,  may  be 
explained  by  such  organs  having  been  gradually  acquired  by  the 
one  sex,  and  then  transmitted  in  a  more  or  less  imperfect  state 
to  the  other.  When  we  treat  of  sexual  selection,  we  shall  meet 
with  innumerable  instances  of  this  form  of  transmission, — as  in 
the  case  of  the  spurs,  plumes,  and  brilliant  colors,  acquired  for 
battle  or  ornament  hy  male  birds,  and  inherited  by  the  females 
in  an  imperfect  or  rudimentary  condition. 

The  possession  by  male  mammals  of  functionally  imperfect 
mammary  organs  is,  in  some  respects,  especially  curious.  The 
Monotremata  have  the  proper  milk-secreting  glands  with  orifices, 
but  no  nipples;  and  as  these  animals  stand  at  the  very  base  of 
the  mammalian  series,  it  is  probable  that  the  progenitors  of 
the  class  also  had  milk-secreting  glands,  but  no  nipples.  This 
conclusion  is  supported  by  what  is  known  of  their  manner  of 
development;    for  Professor  Turner  informs  me,  on  the  authority 

28  This  is  the  conclusion  of  Prof.  Gegenbaur,  one  of  the  highest  au- 
thorities in  comparative  anatomy;  see  'Grundzuge  der  vergleich. 
Anat.'  1870,  s.  876.  The  result  has  been  arrived  at  chiefly  from  the 
study  of  the  Amphibia;  but  it  appears  from  the  researches  of  Wal- 
deyer  (as  quoted  in  'Journal  of  Anat.  and  Phys.'  1869,  p.  161),  that  the 
sexual  organs  of  even  "the  higher  vertebrata  are,  in  their  early  ccndi- 
"tion,  hermaphrodite."  Similar  views  have  long  been  held  by  some 
authors,   though  until  recently  without  a  firm  basis. 

27  The  male  Thylacinus  offers  the  best  instance.  Owen,  'Anatomy  of 
Vertebrates,'  vol.  iii.  p.  771. 

28  Hermaphroditism  has  been  observed  in  several  species  of  Serranus, 
as  well  as  in  some  other  fishes,  where  it  is  either  normal  and  sym- 
metrical, or  abnormal  and  unilateral.  Dr.  Zouteveen  has  given  me  ref- 
erences on  this  subject,more  especially  to  a  paper  by  Prof.Ralbertsma, 
in  the  'Transact,  of  the  Dutch  Acad,  of  Sciences,'  vol.  xvi.  Dr.  Gun- 
ther  doubts  the  fact,  but  it  has  now  been  recorded  by  too  many  good 
observers  to  be  any  longer  disputed.  Dr.  M.  Lessona  writes  to  me, 
that  he  has  verified  the  observations  made  by  Cavolini  on  Serranus. 
Prof.  Ercolani  has  recently  shown  ('Accad.  delle  Scienze,'  Bologna, 
Dec.  28,  1871)  that  eels  are  androgynous. 


158  THE   DESCENT  OF  MAN. 

of  Kolliker  and  Langer,  that  in  the  embryo  the  mammary  glands 
can  be  distinctly  traced  before  the  nipples  are  in  the  least 
visible;  and  the  development  of  successive  parts  in  the  indi- 
vidual generally  represents  and  accords  with  the  development  of 
successive  beings  in  the  same  line  of  descent.  The  Marsupials 
differ  from  the  Monotremata  by  possessing  nipples;  so  that 
probably  these  organs  were  first  acquired  by  the  Marsupials, 
after  they  had  diverged  from,  and  risen  above,  the  Monotremata, 
and  were  then  transmitted  to  the  placental  mammals.-^  No  one 
will  suppose  that  the  Marsupials  still  remained  androgynous  after 
they  had  approximately  acquired  their  present  structure.  How 
then  are  we  to  account  for  male  mammals  possessing  mammae? 
It  is  possible  that  they  were  first  developed  in  the  females  and 
then  transferred  to  the  males;  but  from  what  follows  this  is 
hardly  probable. 

It  may  be  suggested,  as  another  view,  that  long  after  the 
progenitors  of  the  whole  mammalian  class  had  ceased  to  be 
androgynous,  both  sexes  yielded  milk,  and  thus  nourished  their 
young;  and  in  the  case  of  the  Marsupials,  that  both  sexes  carried 
their  young  in  marsupial  sacks.  This  will  not  appear  altogether 
improbable,  if  we  reflect  that  the  males  of  existing  syngnathous 
fishes  receive  the  eggs  of  the  females  in  their  abdominal  pouches, 
hatch  them,  and  afterwards,  as  some  believe,  nourish  the 
young ;^° — that  certain  other  male  fishes  hatch  the  eggs  within 
their  mouths  or  branchial  cavities; — that  certain  male  toads 
take  the  chaplets  of  eggs  from  the  females,  and  wind  them  round 
their  own  thighs,  keeping  them  there  until  the  tadpoles  are 
born; — that  certain  male  birds  undertake  the  whole  duty  of 
incubation,  and  that  male  pigeons,  as  well  as  the  females,  feed 
their  nestlings  with  a  secretion  from  their  crops.  But  the  above 
suggestion  first  occurred  to  me  from  the  mammary  glands  of 
male  mammals  being  so  much  more  perfectly  developed  than 
the  rudiments  of  the  other  accessory  reproductive  parts,  which 
are  found  in  the  one  sex  though  proper  to  the  other.    The  mam- 

2»  Prof.  Gag-enbaur  has  shown  ('Jenaische  Zeitschrift,'  Bd.  vii.  p.  212) 
that  two  distinct  types  of  nipples  prevail  throughout  the  several  mam- 
malian orders,  but  that  it  is  quite  intelligible  how  both  could  have 
been  derived  from  the  nipples  of  the  Marsupials,  and  the  latter  from 
those  of  the  Monotremata.  See,  also,  a  memoir  by  Dr.  Huss,  on 
the  mammary  glands  ibid.  B.  viii.  p.  176. 

^  Mr.  Lockwood  believes  (as  quoted  in  'Quart.  Journal  of  Science,' 
April,  1868,  p.  269),  from  what  he  has  observed  of  the  development  of 
Hippocampus,  that  the  walls  of  the  abdominal  pouch  of  the  male  in 
some  way  afford  nourishment.  On  male  fishes  hatching  the  ova  in 
their  mouths,  see  a  very  interesting  paper  by  Prof.  Wyman,  in  'Proc. 
Boston  Soc.  of  Nat.  Hist.'  Sept.  15,  1857;  also  Prof.  Turner,  in  'Journal 
of  Anat.  and  Phys.'  Nov.  1,  1866,  p.  78.  Dr.  Gunther  has  likewise  de- 
scribed similar  cases. 


APFINITIES    AND    GENEALOGY.  159 

mary  glands  and  nipples,  as  they  exist  in  male  mammals,  can 
indeed  hardly  be  called  rudimentary;  they  are  merely  not  fully 
developed,  and  not  functionally  active.  They  are  sympathetically 
affected  under  the  influence  of  certain  diseases,  like  the  same 
organs  in  the  female.  They  often  secrete  a  few  drops  of  milk  at 
birth  and  at  puberty:  this  latter  fact  occurred  in  the  curious 
case,  before  referred  to,  where  a  young  man  possessed  two  pairs 
of  mammse.  In  man  and  some  other  male  mammals  these  organs 
have  been  known  occasionally  to  become  so  well  developed  dur- 
ing maturity  as  to  yield  a  fair  supply  of  milk.  Now  if  we  sup- 
pose that  during  a  former  prolonged  period  male  mammals  aided 
the  females  in  nursing  their  offspring,-^  and  that  afterwards  from 
some  cause  (as  from  the  production  of  a  smaller  number  of 
young)  the  males  ceased  to  give  this  aid,  disuse  of  the  organs 
during  maturity  would  lead  to  their  becoming  inactive;  and 
from  two  well-known  principles  of  inheritance,  this  state  of 
inactivity  would  probably  be  transmitted  to  the  males  at  the 
corresponding  age  of  maturity.  But  at  an  earlier  age  these  or- 
gans would  be  left  unaffected,  so,  that  they  would  be  almost 
equally  well  developed  in  the  young  of  both  sexes. 

Conclusion. — Von  Baer  has  defined  advancement  or  progress  in 
the  organic  scale  better  than  any  one  else,  as  resting  on  the 
amount  of  differentiation  and  specialization  of  the  several  parts 
of  a  being, — when  arrived  at  maturity;  as  I  should  be  inclined  to 
add.  Now  as  organisms  have  become  slowly  adapted  to  diver- 
sified lines  of  life  by  means  of  natural  selection,  their  parts  will 
have  become  more  and  more  differentiated  and  specialized  for 
various  functions,  from  the  advantage  gained  by  the  division  of 
physiological  labor.  The  same  part  appears  often  to  have  been 
modified  first  for  one  purpose,  and  then  long  afterwards  for  some 
other  and  quite  distinct  purpose;  and  thus  all  the  parts  are 
rendered  more  and  more  complex.  But  each  organism  still  re- 
tains the  general  type  of  structure  of  the  progenitor  from  which 
it  was  aboriginally  derived.  In  accordance  with  this  view  it 
seems,  if  we  turn  to  geological  evidence,  that  organization  on 
the  whole  has  advanced  throughout  the  world  by  slow  and  in- 
terrupted steps.  In  the  great  kingdom  of  the  Vertebrata  it  has 
culminated  in  man.  It  must  not,  however,  be  supposed  that 
groups  of  organic  beings  are  always  supplanted,  and  disappear 
as  soon  as  they  have  given  birth  to  other  and  more  perfect 
groups.  The  latter,  though  victorious  over  their  predecessors, 
may  not  have  become  better  adapted  for  all  places  in  the  economy 
of  nature.    Some  old  forms  appear  to  have  survived  from  inhab- 


31  Madlle.  C.  Royer  has  suggested  a  similar  view  in  her  'Origine  de 
I'Homme,'  &c.,  1870. 


160  THE  DESCENT  OF  MAN. 

iting  protected  sites,  where  they  have  not  been  exposed  to  very 
severe  competition;  and  these  often  aid  us  in  constructing  our 
genealogies,  by  giving  us  a  fair  idea  of  former  and  lost  popula- 
tions. But  we  must  not  fall  into  the  error  of  looking  at  the  exist- 
ing members  of  any  lowly-organized  group  as  perfect  representa- 
tives of  their  ancient  predecessors. 

The  most  ancient  progenitors  in  the  kingdom  of  the  Vertebrata, 
at  which  we  are  able  to  obtain  an  obscure  glance,  apparently 
consisted  of  a  group  of  marine  animals,^-  resembling  the  larvae  of 
existing  Ascidians.  These  animals  probably  gave  rise  to  a  group 
of  fishes,  as  lowly  organized  as  the  lancelet;  and  from  these 
the  Ganoids,  and  other  fishes  like  the  Lepidosiren,  must  have 
been  developed.  From  such  fish  a  very  small  advance  would 
carry  us  on  to  the  Amphibians.  We  have  seen  that  birds  and 
reptiles  were  once  intimately  connected  together;  and  the  Mono- 
tremata  now  connect  mammals  with  reptiles  in  a  slight  degree. 
But  no  one  can  at  present  say  by  what  line  of  descent  the  three 
higher  and  related  classes,  namely,  mammals,  birds,  and  reptiles,, 
were  derived  from  the  two  lower  vertebrate  classes,  namely, 
amphibians  and  fishes.  In  the  class  of  mammals  the  steps  are 
not  difficult  to  conceive  which  led  from  the  ancient  Monotremata 
to  the  ancient  Marsupials;    and  from  these  to  the  early  progeni- 


es The  inhabitants  of  the  seashore  must  be  greatly  affected  by  the 
tides;    animals   living-    either    about   the    mean    high-water   mark,    or 
about  the  mean  low-water  mark,  pass  through  a  complete  cycle  of 
tidal  changes  in  a  fortnight.    Consequently,  their  food  supply  will  un- 
dergo marked  changes  week  by  week.    The  vital   functions  of  such 
animals,    living    under    these   conditions    for    many    generations,    can 
hardly  fail  to  run  their  course  in  regular  weekly  periods.    Now  it  is 
a  mysterious  fact  that  in  the  higher  and  now  terrestrial  Vertebrata, 
as  well  as  in  other  classes,  many  normal  and  abnormal  processes  have 
one  or  more  whole  weeks   as  their  periods;    this  would  be  rendered 
intelligible  if  the  Vertebrata  are  descended  from  an  animal  allied  to 
the  existing  tidal  Ascidians.     Many  instances  of  such  periodic  proc- 
esses might  be  given,  as  the  gestation  of  mammals,   the  duration  of 
fevers,   &c.     The  hatching  of  eggs  affords  also  a  good  example,  for, 
according  to  Mr.   Bartlett  ('Land  and  Water,'  Jan.   7,  1871),   the  eggs 
of  the  pigeon  are  hatched  in  two  weeks;    those  of  the  fowl  in  three; 
those  of  the  duck  in  four;    those  of  the  goose  in  five;    and  those  of  the 
ostrich  in  seven  weeks.    As  far  as  we  can  judge,  a  recurrent  period, 
if  approximately  of  the  right  duration   for  any  process  or  function, 
would  not,    when  once   gained,   be   liable   to   change;    consequently  it 
might  be  thus  transmitted  through  almost  any  number  of  generations. 
But  if  the  function  changed,   the  period  would  have  to  change,   and 
would  be  apt  to  change  almost  abruptly  by  a  whole  week.    This  con- 
clusion, if  sound,  is  highly  remarkable;    for  the  period  of  gestation  in 
each  mammal,  and  the  hatching  of  each  bird's  eggs,  and  many  other 
vital  processes,  thus  betray  to  us  the  primordial  birth-place  of  these 
animals. 


AFFINITIES    AND    GENEALOGY.  161 

tors  of  the  placental  mammals.  We  may  thus  ascend  to  the  Le- 
murid^;  and  the  interval  is  not  very  wide  from  these  to  the 
Simiadae.  The  Simiadse  then  branched  off  into  two  great  stems, 
the  New  World  and  Old  World  monkeys;  and  from  the  latter, 
at  a  remote  period,  Man,  the  wonder  and  glory  of  the  Universe, 
proceeded.  .^ 

Thus  we  have  given  to  man  a  pedigree  of  prodigious  length,  hut  J. 
not,  it  may  be  said,  of  noble  quality.  The  world,  it  has  ofteir*' 
been  remarked,  appears  as  if  it  had  long  been  preparing  for  the 
advent  of  man:  and  this,  in  one  sense  is  strictly  true,  for  he 
owes  his  birth  to  a  long  line  of  progenitors.  If  any  single  link 
in  this  chain  had  never  existed,  man  would  not  have  been  exactly 
what  he  now  is.  Unless  we  wilfully  close  our  eyes,  we  may,  with 
our  present  knowledge,  approximately  recognize  our  parentage; 
nor  need  we  feel  ashamed  of  it.  The  most  humble  organism  is 
something  much  higher  than  the  inorganic  dust  under  our  feet; 
and  no  one  with  an  unbiased  mind  can  study  any  living  creature, 
however  humble,  without  being  struck  with  enthusiasm  at  its 
marvelous  structure  and  properties. 


162  THE  DESCENT  OP  MAN. 


CHAPTER    VII. 
ON  THE  RACES  OF  MAN. 

The  nature  and  value  of  specific  characters— Application  to  the  races 
of  man— Arguments  in  favor  of,  and  opposed  to,  ranking  the  so- 
called  races  of  man  as  distinct  species— Sub-species— Monogenists 
and  polygenists—Convergence  of  character— Numerous  points  of 
resemblance  in  body  and  mind  between  the  most  distinct  races  of 
man— The  state  of  man  when  he  first  spread  over  the  earth — Each 
race  not  descended  from  a  single  pair — The  extinction  of  races — 
The  formation  of  races — The  efCects  of  crossing— Slight  influence 
of  the  direct  action  of  the  conditions  of  life— Slight  or  no  influence 
of  natural  selection— Sexual  selection. 

It  is  not  my  intention  here  to  describe  the  several  so-called 
races  of  men;  but  I  am  about  to  inquire  what  is  the  value  of  the 
differences  between  them  under  a  classificatory  point  of  view, 
and  how  they  have  originated.  In  determining  whether  two  or 
more  allied  forms  ought  to  be  ranked  as  species  or  varieties, 
naturalists  are  practically  guided  by  the  following  considera- 
tions; namely,  the  amount  of  difference  between  them,  and 
whether  such  differences  relate  to  few  or  many  points  of  struc- 
ture, and  whether  they  are  of  physiological  importance;  but 
more  especially  whether  they  are  constant.  Constancy  of  char- 
acter is  what  is  chiefly  valued  and  sought  for  by  naturalists. 
Whenever  it  can  be  shown,  or  rendered  probable,  that  the  forms 
in  question  have  remained  distinct  for  a  long  period,  this  be- 
comes an  argument  of  much  weight  in  favor  of  treating  them  as 
species.  Even  a  slight  degree  of  sterility  between  any  two  forms 
when  first  crossed,  or  in  their  offspring,  is  generally  considered 
as  a  decisive  test  of  their  specific  distinctness;  and  their  con- 
tinued persistence  without  blending  within  the  same  area,  is 
usually  accepted  as  sufficient  evidence,  either  of  some  degree  of 
mutual  sterility,  or  in  the  case  of  animals  of  some  mutual  repug- 
nance to  pairing. 

Independently  of  fusion  from  intercrossing,  the  complete  ab- 
sence, in  a  well-investigated  region,  of  varieties  linking  together 
any  two  closely-allied  forms,  is  probably  the  most  important  of 
all  the  criterions  of  their  specific  distinctness;  and  this  is  a  some* 


THE    RACES    OF    MAN. 

what  different  consideration  from  mere  constancy  of  character, 
for  two  forms  may  be  highly  variable  and  yet  not  yield  inter- 
mediate varieties.  Geographical  distribution  is  often  brought 
into  play  unconsciously  and  sometimes  consciously;  so  that 
forms  living  in  two  widely  separated  areas,  in  which  most  of  the 
other  inhabitants  are  specifically  distinct,  are  themselves  usually 
looked  at  as  distinct;  but  in  truth  this  affords  no  aid  in  dis- 
tinguishing geographical  races  from  so-called  good  or  true  species. 

Now  let  us  apply  these  generally-admitted  principles  to  the 
races  of  man,  viewing  him  in  the  same  spirit  as  a  naturalist  would 
any  other  animal.  In  regard  to  the  amount  of  difference  between 
the  races,  we  must  make  some  allowance  for  cur  nice  powers  of 
discrimination  gained  by  the  long  habit  of  observing  ourselves. 
In  India,  as  Elphinstone  remarks,  although  a  newly-arrived  Eu- 
ropean cannot  at  first  distinguish  the  various  native  races,  yet 
they  soon  appear  to  him  extremely  dissimilar;^  and  the  Hindoo 
cannot  at  first  perceive  any  difference  between  the  several  Eu- 
ropean nations.  Even  the  most  distinct  races  of  man  are  much 
more  like  each  other  in  form  than  would  at  first  be  supposed; 
certain  negro  tribes  must  be  excepted,  whilst  others,  as  Dr.  Rohlfs 
writes  to  me,  and  as  I  have  myself  seen,  have  Caucasian 
features.  This  general  similarity  is  well  shown  by  the  French 
photographs  in  the  Collection  Anthropologique  du  Museum  de 
Paris  of  the  men  belonging  to  various  races,  the  greater  number 
of  which  might  pass  for  Europeans,  as  many  persons  to  whom 
I  have  shown  them  have  remarked.  Nevertheless,  these  men,  if 
seen  alive,  would  undoubtedly  appear  very  distinct,  so  that  we 
are  clearly  much  influenced  in  our  judgment  by  the  mere  color 
of  the  skin  and  hair,  by  slight  differences  in  the  features,  and 
by  expression. 

There  is,  however,  no  doubt  that  the  various  races,  when  care- 
fully compared  and  measured,  differ  much  from  each  other, — as 
in  the  texture  of  the  hair,  the  relative  proportions  of  all  parts 
of  the  body,^  the  capacity  of  the  lungs,  the  form  and  capacity  of 
the  skull,  and  even  in  the  convolutions  of  the  brain."  But  it 
would  be  an  endless  task  to  specify  the  numerous  points  of  dif- 
ference.   The  races  differ  also  in  constitution,  in  acclimatization 

1  'History  of  India,'  1841,  vol.  i.  p.  323.  Father  Ripa  makes  exactly 
the  same  remark  with  respect  to  the  Chinese. 

2  A  vast  number  of  measurements  of  Whites,  Blacks,  and  Indians, 
are  given  in  the  'Investigations  in  the  Militai-y  and  Anthropolog.  Sta- 
tistics of  American  Soldiers,'  by  B.  A.  Gould,  1869,  pp.  298-358;  'On  the 
capacity  of  the  lungs,'  p.  471.  See,  also,  the  numerous  and  valuable 
tables,  by  Dr.  Weisbach,  from  the  observations  of  Dr.  Scherzer  and 
Dr.  Schwarz,  in  the  'Reise  der  Novara:   Anthropolog.  Theil,'  1867. 

2  See,  for  instance,  Mr.  Marshall's  account  of  the  brain  of  a  Bush- 
womaii  in  'PhiL  Transact.'  1864,  p.  519. 


164  THE  DESCENT  OF  MAN. 

and  In  liability  to  certain  diseases.  Their  mental  cliaracteristies 
are  likewise  very  distinct;  chiefly  as  it  would  appear  in  their 
emotional,  but  partly  in  their  intellectual  faculties.  Every  one 
who  has  had  the  opportunity  of  comparison,  must  have  been 
struck  with  the  contrast  between  the  taciturn,  even  morose, 
aborigines  of  S,  America  and  the  light-hearted,  talkative  negroes. 
There  is  a  nearly  similar  contrast  between  the  Malays  and  the 
Papuans,*  who  live  under  the  same  physical  conditions,  and  are 
separated  from  each  other  only  by  a  narrow  space  of  sea. 

We  will  first  consider  the  arguments  which  may  be  advanced 
in  favor  of  classing  the  races  of  man  as  distinct  species,  and 
then  the  arguments  on  the  other  side.  If  a  naturalist,  who  had 
never  before  seen  a  Negro,  Hottentot,  Australian,  or  Mongolian, 
were  to  compare  them,  he  would  at  once  perceive  that  they 
differed  in  a  multitude  of  characters,  some  of  slight  and  some  of 
considerable  importance.  On  inquiry  he  would  find  that  they 
were  adapted  to  live  under  widely  different  climates,  and  that 
they  differed  somewhat  in  bodily  constitution  and  mental  dis- 
position. If  he  were  then  told  that  hundreds  of  similar  specimens 
could  be  brought  from  the  same  countries,  he  would  assuredly 
declare  that  they  were  as  good  species  as  many  to  which  he  had 
been  in  the  habit  of  afiixing  specific  names.  This  conclusion 
would  be  greatly  strengthened  as  soon  as  he  had  ascertained  that 
these  forms  had  all  retained  the  same  character  for  many  cen- 
turies; and  that  negroes,  apparently  identical  with  existing 
negroes,  had  lived  at  least  4000  years  ago.^  He  would  also  hear, 
on  the  authority  of  an  excellent  observer.  Dr.  Lund,^  that  the 


*  Wallace,   'The  Malay  Archipelago,'  vol.  ii.  1869,  p.  178. 

5  With  respect  to  the  figures  in  the  famous  Egyptian  caves  of  Abou- 
Simbel,  M.  Pouchet  says  ('The  Plurality  of  the  Human  Races,'  Eng-. 
translat.  1864,  p.  50),  that  he  was  far  from  finding  recognizable  repre- 
sentations of  the  dozen  or  more  nations  which  some  authors  believ© 
that  they  can  recognize.  Even  some  of  the  most  strongly-marked' 
races  cannot  be  identified  with  that  degree  of  unanimity  which  might 
have  been  expected  from  what  has  been  written  on  the  subject.  Thus 
Messrs.  Nott  and  Gliddon  ('Types  of  Mankind,'  p.  148)  state  that 
Rameses  II.,  or  the  Great,  has  features  superbly  European;  whereas 
Knox,  another  firm  believer  in  the  specific  distinctness  of  the  races 
of  man  ('Races  of  Man,'  1850,  p.  201),  speaking  of  young  Memnon  (the 
same  as  Rameses  II.,  as  I  am  informed  by  Mr.  Birch),  insists  in  the 
strongest  manner  that  he  is  identical  in  character  with  the  Jews  of 
Antwerp.  Again,  when  I  looked  at  the  statute  of  Amunoph  III.,  I 
agreed  with  two  officers  of  the  establishment,  both  competent  judges, 
that  he  had  a  strongly  marked  negro  type  of  features;  but  Messrs. 
Nott  and  Gliddon  (ibid.  p.  146,  fig.  53)  describe  him  as  a  hybrid  but 
not  of  "negro  intermixture." 

«  As  quoted  by  Nott  and  Gliddon,  'Types  of  Mankind,'  1854,  p.  439. 
They  give  also  corroborative  evidence;  trtit  C,  .VTijgt  tMnks  that  tiiQ 
S'ubj'ect  I'e^q'uir^  fHirthtedr  ia«vfegtige*ion. 


TKE    RACES    OF    MAN.  165 

human  skulls  found  in  the  caves  of  Brazil,  entombed  with  many- 
extinct  mammals,  belonged  to  the  same  type  as  that  now  pre- 
vailing throughout  the  American  Continent. 

Our  naturalist  would  then  perhaps  turn  to  geographical  dis- 
tribution, and  he  would  probablj'-  declare  that  those  forms  must 
be  distinct  species,  which  differ  not  only  in  appearance,  but 
are  fitted  for  hot,  as  well  as  damp  or  dry  countries,  and  for  the 
Arctic  regions.  He  might  appeal  to  the  fact  that  no  species  in 
the  group  next  to  man,  namely  the  Quadrumana,  can  resist  a  low 
temperature,  or  any  considerable  change  of  climate;  and  that 
the  species  which  come  nearest  to  man  have  never  been  reared 
to  maturity,  even  under  the  temperate  climate  of  Europe.  He 
would  be  deeply  impressed  with  the  fact,  first  noticed  by  Agassiz,'' 
that  the  different  races  of  man  are  distributed  over  the  world  in 
the  same  zoological  provinces,  as  those  inhabited  by  undoubtedly 
distinct  species  and  genera  of  mammals.  This  is  manifestly  the 
case  with  the  Australian,  Mongolian,  and  Negro  races  of  man;  in 
a  less  well-marked  manner  with  the  Hottentots;  but  plainly  with 
the  Papuans  and  Malays,  who  are  separated,  as  Mr.  Wallace  has 
shown,  by  nearly  the  same  line  which  divides  the  great  Malayan 
and  Australian  zoological  provinces.  The  Aborigines  of  America 
range  throughout  the  Continent;  and  this  at  first  appears  op- 
posed to  the  above  rule,  for  most  of  the  productions  of  the  South- 
ern and  Northern  halves  differ  widely:  yet  some  few  living  forms, 
as  the  opossum,  range  from  the  one  into  the  other,  as  did  formerly 
some  of  the  gigantic  Edentata.  The  Esquimaux,  like  other  Arctic 
animals,  extend  round  the  whole  polar  regions.  It  should  be 
observed  that  the  amount  of  difference  between  the  mammals  of 
the  several  zoological  provinces  does  not  correspond  with  the 
degree  of  separation  between  the  latter;  so  that  it  can  hardly  be 
considered  as  an  anomaly  that  the  Negro  differs  more,  and  the 
American  much  less  from  the  other  races  of  man,  than  do  the 
mammals  of  the  African  and  American  continents  from  the  mam- 
mals of  the  other  provinces.  Man,  it  may  be  added,  does  not 
appear  to  have  aboriginally  inhabited  any  oceanic  island;  and  in 
this  respect  he  resembles  the  other  members  of  his  class. 

In  determining  whether  the  supposed  varieties  of  the  same 
kind  of  domestic  animal  should  be  ranked  as  such,  or  as  specifical- 
ly distinct,  that  is,  whether  any  of  them  are  descended  from 
distinct  wild  species,  every  naturalist  would  lay  much  stress  on 
the  fact  of  their  external  parasites  being  specifically  distinct. 
All  the  m-ore  stress  would  be  laid  on  this  fact,  as  it  would  be  an 
exceptional  one;  for  I  am  informed  by  Mr.  Denny  that  the  most 
different  kinds  of  dogs,  fowls,  and  pigeons,  in  England,  are  in- 


7 'Diversits'  cf  .Origrki  of  this  Human  Races,'  in  thte  "^Ghristia^  Ex- 

12 


16g  THE   DESCENT  OF  MAN. 

fested  by  the  same  species  of  Pediculi  or  lice.  Now  Mr.  A. 
Murray  lias  carefully  examined  the  Pediculi  collected  in  different 
countries  from  the  different  races  of  man;^  and  he  finds  that 
they  differ,  not  only  in  color,  but  in  the  structure  of  their  claws 
and  limbs.  In  every  case  in  which  many  specimens  were  obtained 
the  differences  were  constant.  The  surgeon  of  a  whaling  ship 
in  the  Pacific  assured  me  that  wh^n  the  Pediculi,  with  which 
some  Sandwich  Islanders  on  board  swarmed,  strayed  on  to  the 
bodies  of  the  English  sailors,  they  died  in  the  course  of  three 
or  four  days.  These  Pediculi  were  darker  colored,  and  appeared 
different  from  those  proper  to  the  natives  of  Chiloe  in  South 
America,  of  which  he  gave  me  specimens.  These,  again,  appeared 
larger  and  much  softer  than  European  lice.  Mr.  Murray  pro- 
cured four  kinds  from  Africa,  namely  from  the  Negroes  of  the 
Eastern  and  Western  coasts,  from  the  Hottentots  and  Kafiirs; 
two  kinds  from  the  natives  of  Australia;  two  from  North  and 
two  from  South  America.  In  these  latter  cases  it  may  be  pre- 
sumed that  the  Pediculi  came  from  natives  inhabiting  different 
districts.  With  insects  slight  structural  differences,  if  constant, 
are  generally  esteemed  of  specific  value:  and  the  fact  of  the 
races  of  man  being  infested  by  parasites,  which  appear  to  be 
specifically  distinct,  might  fairly  be  urged  as  an  argument  that 
the  races  themselves  ought  to  be  classed  as  distinct  species. 

Our  supposed  naturalist  having  proceeded  thus  far  in  his  In- 
vestigation, would  next  inquire  whether  the  races  of  men,  when 
crossed,  were  in  any  degree  sterile.  He  might  consult  the  work" 
of  Professor  Broca,  a  cautious  and  philosophical  observer,  and  in 
this  he  would  find  good  evidence  that  some  races  were  quite  fertile 
together,  but  evidence  of  an  opposite  nature  in  regard  to  other 
races.  Thus  it  has  been  asserted  that  the  native  women  of  Aus- 
tralia and  Tasmania  rarely  produce  children  to  European  men; 
the  evidence,  however,  on  this  head  has  now  been  shown  to  be 
almost  valueless.  The  half-castes  are  killed  by  the  pure  blacks: 
and  an  account  has  lately  been  published  of  eleven  half-caste 
youths  murdered  and  burnt  at  the  same  time,  whose  remains 
were  found  by  the  police.^"  Again,  it  has  often  been  said  that 
when  mulattoes  intermarry  they  produce  few  children;    on  the 

8  'Transact.  R.  Soc.  of  Edinburgh,'  vol.  xxii.  1861,  p.  567. 

»  'On  the  Phenomena  of  Hybridity  in  the  Genus  Homo,'  Eng-.  translat. 
1864. 

1°  See  the  interesting  letter  by  Mr.  T.  A.  Murray,  in  the  'Anthropolog. 
Review,'  April,  1868  p.  liii.  In  this  letter  Count  Strzelecki's  statement, 
that  Australian  women  who  have  borne  children  to  a  white  man  are 
afterwards  sterile  with  their  own  race,  is  disproved.  M.  A.  de  Qua' 
trefages  has  also  collected  ('Revue  des  Cours  Scientifiques,'  March, 
1869,.  p.  239)  much  evidence  that  Australians  and  Europeans  are  not 
sterile  when  crossed. 


THE    RACES    OF   MAN.  167 

other  hand,  Dr.  Bachman  of  Charleston"  positively  asserts  that 
he  has  known  mulatto  families  which  have  intermarried  for  sev- 
eral generations,  and  have  continued  on  an  average  as  fertile  as 
either  pure  whites  or  pure  blacks.  Inquiries  formerly  made  by 
Sir  C.  Lyell  on  this  subject  led  him,  as  he  informs  me,  to  the 
same  conciusion.^^  In  the  United  States  the  census  for  the  year 
1854  included,  according  to  Dr.  Bachman,  405,751  mulattoes;  and 
this  number,  considering  all  the  circumstances  of  the  case,  seems 
small;  but  it  may  partly  be  accounted  for  by  the  degraded  and 
anomalous  position  of  the  class,  and  by  the  profligacy  of  the 
women.  A  certain  amount  of  absorption  of  mulattoes  into  ne- 
groes must  always  be  in  progress;  and  this  would  lead  to  an 
apparent  diminution  of  the  former.  The  inferior  vitality  of 
mulattoes  is  spoken  of  in  a  trustworthy  work^^  as  a  well-known 
phenomenon;  and  this,  although  a  different  consideration  from 
their  lessened  fertility,  may  perhaps  be  advanced  as  a  proof  of  the 
specific  distinctness  of  the  parent  races.  No  doubt  both  animal 
and  vegetable  hybrids,  when  produced  from  extremely  distinct 
species,  are  liable  to  premature  death;  but  the  parents  of  mulat- 
toes cannot  be  put  under  the  category  of  extremely  distinct  spe- 
cies. The  common  Mule,  so  notorious  for  long  life  and  vigor, 
and  yet  so  sterile,  shows  how  little  necessary  connection  there  is 
in  hybrids  between  lessened  fertility  and  vitality;  other  analogous 
cases  could  be  cited. 

Even  if  it  should  hereafter  be  proved  that  all  the  races  of  men 
were  perfectly  fertile  together,  he  who  was  inclined  from  other 
reasons  to  rank  them  as  distinct  species,  might  with  justice  argue 
that  fertility  and  sterility  are  not  safe  criterions  of  specific  dis- 
tinctness. We  know  that  these  qualities  are  easily  affected  by 
changed  conditions  of  life,  or  by  close  inter-breeding,  and  that 
they  are  governed  by  highly  complex  laws,  for  instance,  that  of 
the  unequal  fertility  of  converse  crosses  between  the  same  two 
species.  With  forms  which  must  be  ranked  as  undoubted  species, 
a  perfect  series  exists  from  those  which  are  absolutely  sterile 
when  crossed,  to  those  which  are  almost  or  completely  fertile. 

"  'An  Examination  of  Prof.  Agassiz's  Sketch  of  the  Nat.  Provinces 
of  the  Animal  World/  Charleston,  1855,  p.  44. 

12  Dr.  Rohlfs  writes  to  me  that  he  found  the  mixed  races  in  the  Great 
Sahara,  derived  from  Arabs,  Berbers,  and  Neg-roes  of  three  tribes, 
extraordinarily  fertile.  On  the  other  hand,  Mr.  Winwood  Reade  in- 
forms me  that  the  Negroes  on  the  Gold  Coast,  though  admiring  white 
men  and  mulattoes,  have  a  maxim  that  mulattoes  should  not  inter- 
marry, as  the  children  are  few  and  sickly.  This  belief,  as  Mr.  Reade 
remarks,  deserves  attention,  as  white  men  have  visited  and  resided 
on  the  Gold  Coast  for  four  hundred  years,  so  that  the  natives  have  had 
ample  time  to  gain  knowledge  through  experience. 

13  'Military  and  Anthropolog."  Statistics  of  Amejican  Soldiers,'  by 
B.  A.  Gould,  1S69,  p.  319. 


168  THE  DESCENT  OP  MAN. 

The  degrees  of  sterility  do  not  coincide  strictly  with  the  degrees 
of  difference  between  the  parents  in  external  structure  or  habits 
of  life.  Man  in  many  respects  may  be  compared  with  those  ani- 
mals which  have  long  been  domesticated,  and  a  large  body  of 
evidence  can  be  advanced  in  favor  of  the  Pallasian  doctrine,^^ 
that  domestication  tends  to  eliminate  the  sterility  which  is  so 
general  a  result  of  the  crossing  of  species  in  a  state  of  nature. 
From  these  several  considerations,  it  may  be  justly  urged  that 
the  perfect  fertility  of  the  intercrossed  races  of  man,  if  estab- 
lished, would  not  absolutely  preclude  us  from  ranking  them  as 
distinct  species. 

Independently  of  fertility,  the  characters  presented  by  the  off- 
spring from  a  cross  have  been  thought  to  indicate  whether  or  not 
the  parent-forms  ought  to  be  ranked  as  species  or  varieties;  but 
after  carefully  studying  the  evidence,  I  have  come  to  the  con- 
clusion that  no  general  rules  of  this  kind  can  be  trusted.     The 

"  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
p.  109.  I  may  here  remind  the  reader  that  the  sterility  of  species 
when  crossed  is  not  a  specially-acquired  quality,  but,  like  the  inca- 
pacity of  certain  trees  to  be  grafted  tog-ether,  is  incidental  on  oth«r 
acquired  differences.  The  nature  of  these  differences  is  unknown, 
but  they  relate  more  especially  to  the  reproductive  system,  and  much 
less  so  to  external  structure  or  to  ordinary  differences  in  constitution. 
One  important  element  in  the  sterility  of  crossed  species  apparently 
lies  in  one  or  both  having  been  long  habituated  to  fixed  conditions; 
for  we  know  that  changed  conditions  have  a  special  influence  on  the 
reproductive  system,  and  we  have  good  reason  to  believe  (as  before 
remarked)  that  the  fluctuating  conditions  of  domestication  tend  to 
eliminate  that  sterility  which  is  so  general  with  species,  in  a  natural 
state,  when  crossed.  It  has  elsewhere  been  shown  by  me  (ibid.  vol.  ii. 
p.  185,  and  'Origin  of  Species'  5th  edit.,  p.  317),  that  the  sterility  of 
crossed  species  has  not  been  acquired  through  natural  selection:  we 
can  see  that  when  two  forms  have  already  been  rendered  very  sterile, 
it  is  scarcely  possible  that  their  sterility  should  be  augmented  by  the 
preservation  or  survival  of  the  more  and  more  sterile  individuals; 
for  as  the  sterility  increases,  fewer  and  fewer  offspring  will  be  pro- 
duced from  which  to  breed,  and  at  last  only  single  individuals  will 
be  produced  at  the  rarest  intervals.  But  there  is  even  a  higher  grade 
of  sterility  than  this.  Both  Gartner  and  Kolreuter  have  proved  that 
in  genera  of  plants  including  many  species,  a  series  can  be  farmed 
from  species  which  when  crossed  yield  fewer  and  fewer  seeds,  to 
species  which  never  produce  a  single  seed,  but  yet  are  affected  by  the 
pollen  of  the  other  species  as  shown  by  the  swelling  of  the  germen. 
It  is  here  manifestly  impossible  to  select  the  more  sterile  individuals, 
which  have  already  ceased  to  yield  seeds;  so  that  the  acme  of  ster- 
ility, when  the  germen  alone  is  affected  cannot  have  been  gained 
through  selection.  This  acme,  and  no  doubt  the  other  grades  of  ster- 
ility, are  the  incidental  results  of  certain  unknown  differences  in  the 
constitution  oif  the  i^rbUuc'tiVe  ^y^teto  eft  the  ^jneoies  W1a»icfh  arfe 
cro'ss'ed. 


THE    RACES    OF    MAN.  169 

ordinary  result  of  a  cross  is  the  production  of  a  blended  or  inter- 
mediate form;  but  in  certain  cases  some  of  the  oftspiing  take 
closely  after  one  parent-form,  and  some  after  the  other.  This  is 
especially  apt  to  occur  when  the  parents  differ  in  characters 
which  first  appeared  as  sudden  variations  or  monstrosities.^^  I 
refer  to  this  point,  because  Dr.  Rohlfs  informs  me  that  he  has 
frequently  seen  in  Africa  the  offspring  of  negroes  crossed  with 
members  of  other  races,  either  completely  black  or  completely 
white,  or  rarely  piebald.  On  the  other  hand,  it  is  notorious  that 
in  America  mulattoes  commonly  present  an  intermediate  appear- 
ance. 

We  have  now  seen  that  a  naturalist  might  feel  himself  fully 
justified  in  ranking  the  races  of  man  as  distinct  species;  for  he 
has  found  that  they  are  distinguished  by  many  differences  in 
structure  and  constitution,  some  being  of  importance.  These 
differences  have,  also,  remained  nearly  constant  for  very  long 
periods  of  time.  Our  naturalist  will  have  been  in  some  degree 
influenced  by  the  enormous  range  of  man,  which  is  a  great 
anomaly  in  the  class  of  mammals,  if  mankind  be  viewed  as  a 
single  species.  He  will  have  been  struck  with  the  distribution  of 
the  several  so-called  races,  which  accords  with  that  of  other 
undoubtedly  distinct  species  of  mammals.  Finally,  he  might 
urge  that  the  mutual  fertility  of  all  the  races  has  not  as  yet  been 
fully  proved,  and  even  if  proved  would  not  be  an  absolute  proof 
of  their  specific  identity. 

On  the  other  side  of  the  question,  if  our  supposed  naturalist 
were  to  inquire  whether  the  forms  of  man  keep  distinct  like 
ordinary  species,  when  mingled  together  in  large  numbers  in  the 
same  country,  he  would  immediately  discover  that  this  was  by 
no  means  the  case.  In  Brazil  he  would  behold  an  immense  mon- 
grel population  of  Negroes  and  Portuguese;  in  Chiloe,  and  other 
parts  of  South  America,  he  would  behold  the  whole  population 
consisting  of  Indians  and  Spaniards  blended  in  various  degrees." 
In  many  parts  of  the  same  continent  he  would  meet  with  the 
most  complex  crosses  between  Negroes,  Indians,  and  Europeans; 
and  judging  from  the  vegetable  kingdom,  such  triple  crosses 
afford  the  severest  test  of  the  mutual  fertility  of  the  parent-forms. 
In  one  island  of  the  Pacific  he  would  find  a  small  population  of 
mingled  Polynesian  and  English  blood;  and  in  the  Fiji  Archi- 
pelago a  population  of  Polynesian  and  Negritos  crossed  in  all 
degrees.    Many  analogous  cases  could  be  added;    for  instance,  in 


^^  'The  Variation  of  Animals,'  &c.,  vol.  ii.  p.  92. 

18  M.  de  Quatrefages  has  given  ('Anthropolog.  Review,'  Jan.  1869,  p.  22) 
an  interesting  account  of  the  success  and  energy  of  the  Paulistas  in 
Brazil,  who  are  a  much  crossed  race  of  Portuguese  and  Indians,  with 
a  mixture  of  the  blood  of  other  races. 


170  THE   DESCENT  OF  MAN. 

Africa.  Hence  the  races  of  man  are  not  sufficiently  distinct  to 
inhabit  the  same  country  without  fusion;  and  the  absence  of 
fusion  affords  the  usual  and  best  test  of  specific  distinctness. 

Our  naturalist  would  likewise  be  much  disturbed  as  soon  as 
he  perceived  that  the  distinctive  characters  of  all  the  races  were 
highly  variable.  This  fact  strikes  every  one  on  first  beholding 
the  negro  slaves  in  Brazil,  who  have  been  imported  from  all 
parts  of  Africa.  The  same  remark  holds  good  with  the  Poly- 
nesians, and  with  many  other  races.  It  may  be  doubted  whether 
any  character  can  be  named  which  is  distinctive  of  a  race  and  is 
constant.  Savages,  even  within  the  limits  of  the  same  tribe, 
are  not  nearly  so  uniform  in  character,  as  has  been  often  as- 
serted. Hottentot  women  offer  certain  peculiarities,  more  strongly 
marked  than  those  occurring  in  any  other  race,  but  these  are 
known  not  to  be  of  constant  occurrence.  In  the  several  American 
tribes,  color  and  hairiness  differ  considerably;  as  does  color  to  a 
certain  degree,  and  the  shape  of  the  features  greatly  in  the 
Negroes  of  Africa.  The  shape  of  the  skull  varies  much  in  some 
races ;  ^'  and  so  it  is  with  every  other  character.  Now  all  natural- 
ists have  learnt  by  dearly-bought  experience,  how  rash  it  is  to 
attempt  to  define  species  by  the  aid  of  inconstant  characters. 

But  the  most  weighty  of  all  the  arguments  against  treating 
the  races  of  man  as  distinct  species,  is  that  they  graduate  into 
each  other,  independently  in  many  cases,  as  far  as  we  can  judge, 
of  their  having  intercrossed.  Man  has  been  studied  more  carefully 
than  any  other  animal,  and  yet  there  is  the  greatest  possible 
diversity  amongst  capable  judges  whether  he  should  be  classed  as 
a  single  species  or  race,  or  as  two  (Virey),  as  three  (Jacquinot), 
as  four  (Kant),  five  (Blumenbach),  six  (Buff on),  seven  (Hunter), 
eight  (Agassiz),  eleven  (Pickering),  fifteen  (Bory  St.  Vincent), 
sixteen  (Desmoulins),  twenty-two  (Morton),  sixty  (Crawford),  or 
as  sixty-three,  according  to  Burke.^^  This  diversity  of  judgment 
does  not  prove  that  the  races  ought  not  to  be  ranked  as  species, 
but  it  shows  that  they  graduate  into  each  other,  and  that  it  is 
hardly  possible  to  discover  clear  distinctive  characters  between 
them. 

Every  naturalist  who  has  had  the  misfortune  to  undertake  the 
description  of  a  group  of  highly  varying  organisms,  has  en- 
countered cases  (I  speak  after  experience)  precisely  like  that  of 

"  For  instance  with  the  aborigines  of  America  and  Australia.  Prof. 
Huxley  says  ('Transact.  Internat.  Congress  of  Prehist.  Arch.*  1868, 
p.  105)  that  the  skulls  of  many  South  Germans  and  Swiss  are  "as 
"short  and  as  broad  as  those  of  the  Tartars,"  «&c. 

18  See  a  good  discussion  on  this  subject  in  Waitz,  'Introduct.  to  An- 
thropology,' Eng.  translat.  1863,  pp.  19S-20S,  227.  I  have  taken  some  of 
the  above  statements  from  H.  Tuttle's  'Origin  and  Antiquity  of  Physi- 
cal Man,'  Boston,  1866,  p.  35. 


THE    RACES    OF    MAN.  171 

man,  and  if  of  a  cautious  disposition,  he  will  end  by  uniting  all 
the  forms  which  graduate  into  each  other,  under  a  single  species; 
for  he  will  say  to  himself  that  he  has  no  right  to  give  names  to 
objects  which  he  cannot  define.  Cases  of  this  kind  occur  in  the 
Order  which  includes  man,  namely  in  certain  genera  of  monkeys; 
whilst  in  other  genera,  as  in  Cercopithecus,  most  of  the  species 
can  be  determined  with  certainty.  In  the  American  genus  Cebus, 
the  various  forms  are  ranked  by  some  naturalists  as  species,  by 
others  as  mere  geographical  races.  Now  if  numerous  specimens 
of  Cebus  were  collected  from  all  parts  of  South  America,  and 
those  forms  which  at  present  appear  to  be  specifically  distinct, 
were  found  to  graduate  into  each  other  by  close  steps,  they 
would  usually  be  ranked  as  mere  varieties  or  races;  and  this 
course  has  been  followed  by  most  naturalists  with  respect  to  the 
races  of  man.  Nevertheless,  it  must  be  confessed  that  there  are 
forms,  at  least  in  the  vegetable  kingdom,"  which  we  cannot  avoid 
naming  as  species,  but  which  are  connected  together  by  num- 
berless gradations,  independently  of  intercrossing. 

Some  naturalists  have  lately  employed  the  term  "sub-species'* 
to  designate  forms  which  possess  many  of  the  characteristics  of 
true  species,  but  which  hardly  deserve  so  high  a  rank.  Now  if 
we  refiect  on  the  weighty  arguments  above  given,  for  raising  the 
races  of  man  to  the  dignity  of  species,  and  the  insuperable  diffi- 
culties on  the  other  side  in  defining  them,  it  seems  that  the  term 
"sub-species"  might  here  be  used  with  propriety.  But  from 
long  habit  the  term  "race"  will  perhaps  always  be  employed. 
The  choice  of  terms  is  only  so  far  important  in  that  it  is  desirable 
to  use,  as  far  as  possible,  the  same  terms  for  the  same  degrees  of 
difference.  Unfortunately  this  can  rarely  be  done:  for  the  larger 
genera  generally  include  closely-allied  forms,  which  can  be  dis- 
tinguished only  with  much  difficulty,  whilst  the  smaller  genera 
within  the  same  family  include  forms  that  are  perfectly  distinct; 
yet  all  must  be  ranked  equally  as  species.  So  again,  species 
within  the  same  large  genus  by  no  means  resemble  each  other  to 
the  same  degree:  on  the  contrary,  some  of  them  can  generally 
be  arranged  in  little  groups  round  other  species  like  satellites 
round  planets.^" 

The  question  whether  mankind  consists  of  one  or  several  spe- 
cies has  of  late  years  been  much  discussed  by  anthropologists, 
who  are  divided  into  the  two  schools  of  monogenists  and  poly- 
genists.    Those  who  do  not  admit  the  principle  of  evolution,  must 

1®  Prof.  Nageli  has  carefully  described  several  striking  cases  in  his 
'Botanische  Mittheilungen/  B.  ii.  186b",  s.  294-369.  Prof.  Asa  Gray  has 
made  analogous  remarks  on  some  intermediate  forms  in  the  Com- 
positae  of  N.  America. 

20  'Origin  of  Species,'  5th  edit.  p.  68. 


172  THE  DESCENT  OF  MAN. 

look  at  species  as  separate  creations,  or  as  in  some  manner 
as  distinct  entities;  and  they  must  decide  what  forms  of  man 
they  will  consider  as  species  by  the  analogy  of  the  method  com- 
monly pursued  in  ranking  other  organic  beings  as  species.  But  it 
is  a  hopeless  endeavor  to  decide  this  point,  until  some  definition 
of  the  term  "species"  is  generally  accepted;  and  the  definition 
must  not  include  an  indeterminate  element  such  as  an  act  of 
creation.  We  might  as  well  attempt  without  any  definition  to 
decide  whether  a  certain  number  of  houses  should  be  called  a 
village,  town,  or  city.  We  have  a  practical  illustration  of  the 
difficulty  in  the  never-ending  doubts  whether  many  closely  allied 
mammals,  birds,  insects,  and  plants,  which  represent  each  other 
respectively  in  North  America  and  Europe,  should  be  ranked  as 
species  or  geographical  races;  and  the  like  holds  true  of  the  pro- 
ductions of  many  islands  situated  at  some  little  distance  from  the 
nearest  continent. 

Those  naturalists,  on  the  other  hand,  who  admit  the  principle 
of  evolution,  and  this  is  now  admitted  by  the  majority  of  rising 
men,  will  feel  no  doubt  that  all  the  races  of  man  are  descended 
from  a  single  primitive  stock;  whether  or  not  they  may  think 
fit  to  designate  the  races  as  distinct  species,  for  the  sake  of  ex- 
pressing their  amount  of  difference.-^  With  our  domestic  ani- 
mals the  question  whether  the  various  races  have  arisen  from 
one  or  more  species  is  somewhat  different.  Although  it  may  be 
admitted  that  all  the  races,  as  well  as  all  the  natural  species 
within  the  same  genus,  have  sprung  from  the  same  primitive 
stock,  yet  it  is  a  fit  subject  for  discussion,  whether  all  the  do- 
mestic races  of  the  dog,  for  instance,  having  acquired  their  present 
amount  of  difference  since  some  one  species  was  first  domesticated 
by  man;  or  whether  they  owe  some  of  their  characters  to  in- 
heritance from  distinct  species,  which  had  already  been  differen- 
tiated in  a  state  of  nature.  With  man  no  such  question  can  arise, 
for  he  cannot  be  said  to  have  been  domesticated  at  any  particular 
period. 

During  an  early  stage  in  the  divergence  of  the  races  of  man 
from  a  common  stock,  the  differences  between  the  races  and 
their  number  must  have  been  small;  consequently  as  far  as 
their  distinguishing  characters  are  concerned,  they  then  had  less 
claim  to  rank  as  distinct  species  than  the  existing  so-called  races. 
Nevertheless,  so  arbitrary  is  the  term  of  species,  that  such  early 
races  would  perhaps  have  been  ranked  by  some  naturalists  as 
distinct  species,  if  their  differences,  although  extremely  slight, 
had  been  more  constant  than  they  are  at  present,  and  had  not 
graduated  into  each  other. 

21  See  Prof.  Huxley  to  this  effect  in  the  'Fortnightly  Review,'  1865,  p. 
275. 


THE    RACES    OF   MAN.  173 

It  is  however  possible,  though  far  more  probable,  that  the 
early  progenitors  of  man  might  formerly  have  diverged  much  in 
character,  until  they  became  more  unlike  each  other  than  any 
now  existing  races;  but  that  subsequently,  as  suggested  by 
Vogt,^-  they  converged  in  character.  When  man  selects  the  off- 
spring of  two  distinct  species  for  the  same  object,  he  sometimes 
induces  a  considerable  amount  of  convergence,  as  far  as  general 
appearance  is  concerned.  This  is  the  case,  as  shown  by  Von  Nath- 
usius,"^  with  the  improved  breeds  of  the  pig,  which  are  descend- 
ed from  two  distinct  species;  and  in  a  less  marked  manner  with 
the  improved  breeds  of  cattle.  A  great  anatomist,  Gratiolet, 
maintains  that  the  anthropomorphous  apes  do  not  form  a  natural 
sub-group;  but  that  the  orang  is  a  highly  developed  gibbon  or 
semnopithecus,  the  chimpanzee  a  highly  developed  macacus,  and 
the  gorilla  a  highly  developed  mandrill.  If  this  conclusion, 
which  rests  almost  exclusively  on  brain-characters,  be  admitted, 
we  should  have  a  case  of  convergence  at  least  in  external  char- 
acters, for  the  anthropomorphous  apes  are  certainly  more  like 
each  other  in  many  points,  than  they  are  to  other  apes.  All 
analogical  resemblances,  as  of  a  whale  to  a  fish,  may  indeed  be 
said  to  be  cases  of  convergence;  but  this  term  has  never  been  ap- 
plied to  superficial  and  adaptive  resemblances.  It  would,  how- 
ever, be  extremely  rash  to  attribute  to  convergence  close  similar- 
ity of  character  in  many  points  of  structure  amongst  the  modified 
descendants  of  widely  distinct  beings.  The  form  of  'a  crystal  is 
determined  solely  by  the  molecular  forces,  and  it  is  not  surprising 
that  dissimilar  substances  should  sometimes  assume  the  same 
form;  but  with  organic  beings  we  should  bear  in  mind  that  the 
form  of  each  depends  on  an  infinity  of  complex  relations,  namely 
on  variations,  due  to  causes  far  too  intricate  to  be  followed, — 
on  the  nature  of  the  variations  preserved,  these  depending  on  the 
physical  conditions,  and  still  more  on  the  surrounding  organisms 
which  compete  with  each, — and  lastly,  on  inheritance  (in  itself  a 
fluctuating  element)  from  innumerable  progenitors,  all  of  which 
have  had  their  forms  determined  through  equally  complex  rela- 
tions. It  appears  incredible  that  the  modified  descendants  of  two 
organisms,  if  these  differed  from  each  other  in  a  marked  manner, 
should  ever  afterwards  converge  so  closely  as  to  lead  to  a  near 
approach  to  identity  throughout  their  whole  organization  In 
the  case  of  the  convergent  races  of  pigs  above  referred  to,  evi- 
dence of  their  descent  from  two  primitive  stocks  is,  according  to 
Von  Nathusius,  still  plainly  retained,  in  certain  bones  of  their 

22  'Lectures  on  Man,'  Eng-.  translat.  1864,  p.  468. 

2^  'Die  Racen  des  Schweines,'  1860,  s.  46.  'Vorstudien  fur  Geschichte, 
&c.,  Schweineschadel,'  1864,  s.  10^.  With  respect  to  cattle,  see  M.  de 
Quatrefages,  'Unite  de  I'Espece  Humaine,'  1861,  p.  119. 


174  THE   DESCENT  OF  MAN. 

skulls.  If  the  races  of  man  had  descended,  as  Is  supposed  by 
some  naturalists,  from  two  or  more  species,  which  differed  from 
each  other  as  much,  or  nearly  as  much,  as  does  the  orang  from 
the  gorilla,  it  can  hardly  be  doubted  that  marked  differences  in 
the  structure  of  certain  bones  would  still  be  discoverable  in  man 
as  he  now  exists. 


Although  the  existing  races  of  man  differ  in  many  respects,  as 
in  color,  hair,  shape  of  skull,  proportions  of  the  body,  &c.,  yet 
if  their  whole  structure  be  taken  into  consideration  they  are 
found  to  resemble  each  other  closely  in  a  multitude  of  points. 
Many  of  these  are  so  unimportant  or  of  so  singular  a  nature, 
that  it  is  extremely  improbable  that  they  should  have  been  inde- 
pendently acquired  by  aboriginally  distinct  species  or  races.  The 
same  remark  holds  good  with  equal  or  greater  force  with  respect 
to  the  numerous  points  of  mental  similarity  between  the  most 
distinct  races  of  man.  The  American  aborigines,  Negroes  and 
Europeans  are  as  different  from  each  other  in  mind  as  any  three 
races  that  can  be  named;  yet  I  was  incessantly  struck,  whilst 
living  with  the  Fuegians  on  board  the  "Beagle,"  with  the  many 
little  traits  of  character,  showing  how  similar  their  minds  were 
to  ours;  and  so  it  was  with  a  full-blooded  negro  with  whom  I 
happened  once  to  be  intimate. 

He  who  will  read  Mr.  Tylor's  and  Sir  J.  Lubbock's  interesting 
works-*  can  hardly  fail  to  be  deeply  impressed  with  the  close 
similarity  between  the  men  of  all  races  in  tastes,  dispositions  and 
habits.  This  is  shown  by  the  pleasure  which  they  all  take  in 
dancing,  rude  music,  acting,  painting,  tattooing,  and  otherwise 
decorating  themselves;  in  their  mutual  comprehension  of  ges- 
ture-language, by  the  same  expression  in  their  features,  and  by 
the  same  inarticulate  cries,  when  excited  by  the  same  emotions. 
This  similarity,  or  rather  identity,  is  striking,  when  contrasted 
with  the  different  expressions  and  cries  made  by  distinct  species 
of  monkeys.  There  is  good  evidence  that  the  art  of  shooting 
with  bows  and  arrows  has  not  been  handed  down  from  any 
common  progenitor  of  mankind,  yet  as  Westropp  and  Nilsson 
have  remarked,-'  the  stone  arrow-heads,  brought  from  the  most 
distant  parts  of  the  world,  and  manufactured  at  the  most  remote 
periods,  are  almost  identical;  and  this  fact  can  only  be  accounted 
for   by   the   various  races   having   similar   inventive  or  mental 

21  Tylor's  'Early  History  of  Mankind,*  1865;  with  respect  to  gesture- 
language,  see  p.  54.  Lubbock's  'Prehistoric  Times,'  2nd  edit.  1869. 

25  'On  Analogous  Forms  of  Implements,'  in  'Memoirs  of  Anthropolog. 
Soc.,'  by  H.  M.  Westropp.  'The  Primitive  Inhabitants  of  Scandi- 
navia,' Eng.  translat.  edited  by  Sir  J.  Lubbock,  1868,  p.  104. 


THE  RACES  OF  MAN.  175 

powers.  The  same  observation  has  been  made  by  archgeologists" 
with  respect  to  certain  widely-prevalent  ornaments,  such  as  zig- 
zags, &c.;  and  with  respect  to  various  simple  beliefs  and  cus- 
toms, such  as  the  burying  of  the  dead  under  megaiithic  struc- 
tures. I  remember  observing  in  South  America,-'  that  there,  as 
in  so  many  other  parts  of  the  world,  men  have  generally  chosen 
the  summits  of  lofty  hills,  to  throw  up  piles  of  stones,  either  as 
a  record  of  some  remarkable  event,  or  for  burying  their  dead. 

Now  when  naturalists  observe  a  close  agreement  in  numerous 
small  details  of  habits,  tastes,  and  dispositions  between  two  or 
more  domestic  races,  or  between  nearly-allied  natural  forms, 
they  use  this  fact  as  an  argument  that  they  are  descended  from  a 
common  progenitor  who  was  thus  endowed;  and  consequently 
that  all  should  be  classed  under  the  same  species.  The  same 
argument  may  be  applied  with  much  force  to  the  races  of  man. 

As  it  is  improbable  that  the  numerous  and  unimportant  points 
of  resemblance  between  the  several  races  of  man  in  bodily  struc- 
ture and  mental  faculties  (I  do  not  here  refer  to  similar  customs) 
should  all  have  been  independently  acquired,  they  must  have 
been  inherited  from  progenitors  who  had  these  same  characters. 
We  thus  gain  some  insight  into  the  early  state  of  man,  before  he 
had  spread  step  by  step  over  the  face  of  the  earth.  The  spreading 
of  man  to  regions  widely  separated  by  the  sea,  no  doubt,  pre- 
ceded any  great  amount  of  divergence  of  character  in  the  several 
races;  for  otherwise  we  .should  sometimes  meet  with  the  same 
race  in  distinct  continents;  and  this  is  never  the  case.  Sir  J. 
Lubbock,  after  comparing  the  arts  now  practiced  by  savages  iu 
all  parts  of  the  world,  specifies  those  which  man  could  not  have 
known,  when  he  first  wandered  from  his  original  birthplace; 
for  if  once  learnt  they  would  never  have  been  forgotten.^^  He 
thus  shows  that  "the  spear,  which  is  but  a  development  of  the 
"knife-point,  and  the  club,  which  is  but  a  long  hammer,  are  the 
"only  things  left."  He  admits,  however,  that  the  art  of  making 
fire  probably  had  been  already  discovered,  for  it  is  common  to 
all  the  races  now  existing,  and  was  known  to  the  ancient  cave- 
inhabitants  of  Europe.  Perhaps  the  art  of  making  rude  canoes 
or  rafts  was  likewise  known;  but  as  man  existed  at  a  remote 
epoch,  when  the  land  in  many  places  stood  at  a  very  different 
level  to  what  it  does  now,  he  would  have  been  able,  without  the 
aid  of  canoes,  to  have  spread  widely.  Sir  J.  Lubbock  further 
remarks  how  improbable  it  is  that  our  earliest  ancestors  could 
have  "counted  as  high  as  ten,  considering  that  so  many  races 

26  Westropp,    'On  Cromlechs,*  &c.,   'Journal  of  Ethnological  Soc'  as 
given  in  'Scientific  Opinion,'  June  2nd,   1869,   p.  3. 
^  'Journal  of  Researches:  Voyage  of  the  "Beagle,"  '  p.  46. 
28  'Prehistoric  Times,'  1869,  p.  574. 


170  THE  DESCENT  OF  MAN. 

"now  in  existence  cannot  get  beyond  four."  Nevertheless,  at 
this  early  period,  the  intellectual  and  social  faculties  of  man 
could  hardly  have  been  inferior  in  any  extreme  degree  to  those 
possessed  at  present  by  the  lowest  savages;  otherwise  primeval 
man  could  not  have  been  so  eminently  successful  in  the  struggle 
for  life,  as  proved  by  his  early  and  wide  diffusion. 

From  the  fundamental  differences  between  certain  languages, 
some  philologists  have  inferred  that  when  man  first  became 
widely  diffused,  he  was  not  a  speaking  animal;  but  it  may  be 
suspected  that  languages,  far  less  perfect  than  any  now  spoken, 
aided  by  gestures,  might  have  been  used,  and  yet  have  left  no 
traces  on  subsequent  and  more  highly-developed  tongues.  With- 
out the  use  of  some  language,  however  imperfect,  it  appears 
doubtful  whether  man's  intellect  could  have  risen  to  the  standard 
implied  by  his  dominant  position  at  an  early  period. 

Whether  primeval  man,  when  he  possessed  but  few  arts,  and 
those  of  the  rudest  kind,  and  when  his  power  of  language  was 
extremely  imperfect,  would  have  deserved  to  be  called  man,  must 
depend  on  the  definition  which  we  employ.  In  a  series  of  forms 
graduating  insensibly  from  some  ape-like  creature  to  man  as  he 
now  exists,  it  would  be  impossible  to  fix  on  any  definite  point 
when  the  term  "man"  ought  to  be  used.  But  this  is  a  matter  of 
very  little  importance.  So  again,  it  is  almost  a  matter  of  indif- 
ference whether  the  so-called  races  of  man  are  thus  designated, 
or  are  ranked  as  species  or  sub-species;  but  the  latter  term  ap- 
pears the  more  appropriate.  Finally,  we  may  conclude  that  when 
the  principle  of  evolution  is  generally  accepted,  as  it  surely  will 
be  before  long,  the  dispute  between  the  monogenists  and  the 
polygenists  will  die  a  silent  and  unobserved  death. 

One  other  question  ought  not  to  be  passed  over  without  notice, 
namely,  whether,  as  is  sometimes  assumed,  each  sub-species  or 
race  of  man  has  sprung  from  a  single  pair  of  progenitors.  With 
our  domestic  animals  a  new  race  can  readily  be  formed  by  care- 
fully matching  the  varying  offspring  from  a  single  pair,  or  even 
from  a  single  individual  possessing  some  new  character;  but 
most  of  our  races  have  been  formed,  not  intentionally  from  a 
selected  pair,  but  unconsciously  by  the  preservation  of  many  in- 
dividuals which  have  varied,  however  slightly,  in  some  useful  or 
desired  manner.  If  in  one  country  stronger  and  heavier  horses, 
and  in  another  country  lighter  and  fleeter  ones,  were  habitually 
preferred,  we  may  feel  sure  that  two  distinct  sub-breeds  would 
be  produced  in  the  course  of  time,  without  any  one  pair  having 
been  separated  and  bred  from,  in  either  country.  Many  races 
have  been  thus  formed,  and  their  manner  of  formation  is  closely 
analogous  to  that  of  natural  species.  We  know,  also,  that  the 
horses  taken  to  the  Falkland  Islands  have,   during  successive 


THE    EXTINCTION    OF   RACES.  177 

generations,  become  smaller  and  weaker,  whilst  those  which  have 
run  wild  on  the  Pampas  have  acquired  larger  and  coarser  heads; 
and  such  changes  are  manifestly  due,  not  to  any  one  pair,  but  to 
{ill  the  individuals  having  been  subjected  to  the  same  conditions, 
aided,  perhaps,  by  the  principle  of  reversion.  The  new  sub- 
breeds  in  such  cases  are  not  descended  from  any  single  pair,  but 
from  many  individuals  which  have  varied  in  different  degrees, 
but  in  the  same  general  manner;  and  we  may  conclude  that  the 
races  of  man  have  been  similarly  produced,  the  modifications 
being  either  the  direct  result  of  exposure  to  different  conditions, 
or  the  indirect  result  of  some  form  of  selection.  But  to  this 
latter  subject  we  shall  presently  return. 

On  the  Exti7iction  of  the  Races  of  Man. — The  partial  or  com- 
plete extinction  of  many  races  and  sub-races  of  man  is  historically 
known.  Humboldt  saw  in  South  America  a  parrot  which  was 
the  sole  living  creature  that  could  speak  a  word  of  the  language 
of  a  lost  tribe.  Ancient  monuments  and  stone  implements  found 
in  all  parts  of  the  world,  about  which  no  tradition  has  been 
preserved  by  the  present  inhabitants,  indicate  much  extinction. 
Some  small  and  broken  tribes,  remnants  of  former  races,  still 
survive  in  isolated  and  generally  mountainous  districts.  In 
Europe  the  ancient  races  were  all,  according  to  Schaaffhausen,^ 
"lower  in  the  scale  than  the  rudest  living  savages;"  they  must 
therefore  have  differed,  to  a  certain  extent,  from  any  existing 
race.  The  remains  described  by  Professor  Broca  from  Les  Eyzies, 
though  they  unfortunately  appear  to  have  belonged  to  a  single 
family,  indicate  a  race  with  a  most  singular  combination  of  low 
or  simious,  and  of  high  characteristics.  This  race  is  "entirely 
"different  from  any  other,  ancient  or  modern,  that  we  have  ever 
"heard  of."'''  It  differed,  therefore,  from  the  quaternary  race  of 
the  caverns  of  Belgium. 

Man  can  long  resist  conditions  which  appear  extremely  un- 
favorable for  his  existence.^^  He  has  long  lived  in  the  extreme 
regions  of  the  "North,  with  no  wood  for  his  canoes  or  imple- 
ments, and  with  only  blubber  as  fuel,  and  melted  snow  as  drink. 
In  the  southern  extremity  of  America  the  Fuegians  survive  with- 
out the  protection  of  clothes,  or  of  any  building  worthy  to  be 
called  a  hovel.  In  South  Africa  the  aborigines  wander  over  arid 
plains,  where  dangerous  beasts  abound.  Man  can  withstand  the 
deadly  influence  of  the  Terai  at  the  foot  of  the  Himalaya,  and 
the  pestilential  shores  of  tropical  Africa. 

2»  Translation  in  'Anthropolog-ical   Review,'   Oct.  1868,   p.  431. 

^  'Transact.  Internal.  Congress  of  Prehistoric  Arch,'  1S68,  pp.  172-175. 
See  also  Broca  (translation)  in  'Anthropolog-ical  Review,'  Oct.  1868,  p. 
410. 

^  Dr.  Gerland  'Ueber  das  Aussterben  der  Naturvolker,'  1868,  s.  82. 
1? 


178  THE   DESCENT  OF  MAN. 

Extinction  follows  chiefly  from  the  competition  of  tribe  with 
tribe,  and  race  with  race.  Various  checks  are  always  in  action, 
serving  to  keep  dov/n  the  numbers  of  each  savage  tribe, — such 
as  periodical  famines,  nomadic  habits  and  the  consequent  deaths 
of  infants,  prolonged  suckling,  wars,  accidents,  sickness,  licen- 
tiousness, the  stealing  of  women,  infanticide,  and  especially  les- 
sened fertility.  If  any  one  of  these  checks  increases  in  power, 
even  slightly,  the  tribe  thus  affected  tends  to  decrease;  and 
when  of  two  adjoining  tribes  one  becomes  less  numerous  and  less 
powerful  than  the  other,  the  contest  is  soon  settled  by  war, 
slaughter,  cannibalism,  slavery,  and  absorption.  Even  when  a 
weaker  tribe  is  not  thus  abruptly  swept  away,  if  it  once  begins 
to  decrease,  it  generally  goes  on  decreasing  until  it  becomes 
extinct.^- 
~f  When  civilized  nations  come  into  contact  with  barbarians  the 
V  struggle  is  short,  except  where  a  deadly  climate  gives  its  aid  to 
the  native  race.  Of  the  causes  which  lead  to  the  victory  of 
civilized  nations,  some  are  plain  and  simple,  others  complex  and 
obscure.  We  can  see  that  the  cultivation  of  the  land  will  be 
fatal  in  many  ways  to  savages,  for  they  cannot,  or  will  not, 
change  their  habits.  New  diseases  and  vices  have  in  some  cases 
proved  highly  destructive;  and  it  appears  that  a  new  disease 
often  causes  much  death,  until  those  who  are  most  susceptible 
to  its  destructive  influence  are  gradually  weeded  out;^  and  so  it 
may  be  with  the  evil  effects  from  spirituous  liquors,  as  well  as 
with  the  unconquerably  strong  taste  for  them  shown  by  so  many 
savages.  It  further  appears,  mysterious  as  is  the  fact,  that  the 
first  meeting  of  distinct  and  separated  people  generates  disease.^* 
Mr.  Sproat,  who  in  Vancouver  Island  closely  attended  to  the 
subject  of  extinction,  believed  that  changed  habits  of  life,  conse- 
quent on  the  advent  of  Europeans,  induces  much  ill  health.  He 
lays,  also,  great  stress  on  the  apparently  trifling  cause  that  the 
natives  become  "bewildered  and  dull  by  the  new  life  around  them; 
"they  lose  the  motives  for  exertion,  and  get  no  new  ones  in  their 
"place."^^ 

The  grade  of  their  civilization  seems  to  be  a  most  important 
element  in  the  success  of  competing  nations,  A  few  centuries 
ago  Europe  feared  the  inroads  of  Eastern  barbarians;  now  any 
such  fear  w^ould  be  ridiculous.    It  is  a  more  curious  fact,  as  Mr. 

32  Gerland  (ibid.  s.  12)  gives  facts  in  support  of  this  statement. 

^  See  remarks  to  this  effect  in  Sir  H.  Holland's  'Medical  Notes  and 
Reflections,'  1839,  p.  390. 

3*  I  have  collected  ('Journal  of  Researches,  Voyag-e  of  the  "Beagle,"  ' 
p.  435)  a  good  many  cases  bearing  on  this  subject:  see  also  Gerland, 
ibid.  s.  8.  Poeppig  speaks  of  the  "breath  of  civilization  as  poisonous 
*'to  savages." 

^  Sproat,  'Scenes  and  Studies  of  Savage  Life,'  1868,  p.  284. 


THE    EXTINCTION   OF    RACES.  179 

Bagehot  has  remarked,  that  savages  did  not  formerly  waste  away 
before  the  classical  nations,  as  they  now  do  before  modern  civil- 
ized nations;  had  they  done  so,  the  old  moralists  would  have 
mused  over  the  event;  but  there  is  no  lament  in  any  writer  of 
that  period  over  the  perishing  barbarians.^®  The  most  potent  of 
all  the  causes  of  extinction,  appears  in  many  cases  to  be  lessened 
fertility  and  ill-health,  especially  amongst  the  children,  arising 
from  changed  conditions  of  life,  notwithstanding  that  the  nev/ 
conditions  may  not  be  injurious  in  themselves.  I  am  much  in- 
debted to  Mr.  H.  H.  Howorth  for  having  called  my  attention  to 
this  subject,  and  for  having  given  me  information  respecting  it. 
I  have  collected  the  following  cases. 

When  Tasmania  was  first  colonized  the  natives  were  roughly 
estimated  by  some  at  7000  and  by  others  at  20,000.  Their  number 
was  soon  greatly  reduced,  chiefly  by  fighting  with  the  English 
and  with  each  other.  After  the  famous  hunt  by  all  the  colonists, 
when  the  remaining  natives  delivered  themselves  up  to  the  gov- 
ernment, they  consisted  only  of  120  individuals,^"  who  were  in  1832 
transported  to  Flinders  Island.  This  island,  situated  between 
Tasmania  and  Australia,  is  forty  miles  long,  and  from  twelve  to 
eighteen  miles  broad:  it  seems  healthy,  and  the  natives  were 
well  treated.  Nevertheless,  they  suffered  greatly  in  health.  In 
1834  they  consisted  (Bonwick,  p.  250)  of  forty-seven  adult  males, 
forty-eight  adult  females,  and  sixteen  children,  or  in  all  of  111 
souls.  In  1835  only  one  hundred  were  left.  As  they  continued 
rapidly  to  decrease,  and  as  they  themselves  thought  that  they 
should  not  perish  so  quickly  elsewhere,  they  were  removed  in 
1847  to  Oyster  Cove  in  the  southern  part  of  Tasmania.  They  then 
consisted  (Dec.  20th,  1847)  of  fourteen  men,  twenty-two  women 
and  ten  children. ^^  But  the  change  of  site  did  no  good.  Disease 
and  death  still  pursued  them,  and  in  1864  one  man  (who  died  in 
1869),  and  three  elderly  women  alone  survived.  The  infertility 
of  the  w^omen  is  even  a  more  remarkable  fact  than  the  liability 
of  all  to  ill-health  and  death.  At  the  time  when  only  nine  women 
were  left  at  Oyster  Cove,  they  told  Mr.  Bonwick  (p.  386),  that  only 
two  had  ever  borne  children:  and  these  two  had  together  pro- 
duced only  three  children! 

With  respect  to  the  cause  of  this  extraordinary  state  of  things, 
Dr.  Story  remarks  that  death  followed  the  attempts  to  civilize 
the  natives.  "If  left  to  themselves  to  roam  as  they  were  wont 
"and  undisturbed,  they  would  have  reared  more  children,  and 

38  Bagehot,  'Physics  and  Politics,'  'Fortnightly  Review,'  April  1,  1868, 
p.  455. 

37  All  the  statements  here  given  are  taken  from  'The  last  of  the  Tas- 
manians,'  by  J.  Bonwick,  1870. 

38  This  is  the  statement  of  the  Governor  of  Tasmania,  Sir  W.  Deni- 
son,  'Varieties  of  Vice-Regal  Life,'  1S70,  vol.  i.  p.  67. 


180  THE  DESCENT  OF  MAN. 

"there  would  have  been  less  mortality."  Another  careful  ob- 
server of  the  natives,  Mr.  Davis,  remarks,  "The  births  have  been 
"few  and  the  deaths  numerous.  This  may  have  been  in  a  great 
"measure  owing  to  their  change  of  living  and  food;  but  more  so 
"to  their  banishment  from  the  mainland  of  Van  Diemen's  Land, 
"and  consequent  depression  of  spirits"  (Bonwick,  pp.  388,  390). 

Similar  facts  have  been  observed  in  two  widely  difterent  parts 
of  Australia.  The  celebrated  explorer,  Mr.  Gregory,  told  Mr. 
Bonwick,  that  in  Queensland  "the  want  of  reproduction  was  be- 
"ing  already  felt  with  the  blacks,  even  in  the  most  recently  settled 
"parts,  and  that  decay  would  set  in."  Of  thirteen  aborigines  from 
Shark's  Bay  who  visited  Murchison  River,  twelve  died  of  con- 
sumption within  three  months.^^ 

The  decrease  of  the  Maories  of  New  Zealand  has  been  caj'efully 
investigated  by  Mr.  Fenton,  in  an  admirable  report,  from  which 
all  the  following  statements,  with  one  exception,  are  taken.^" 
The  decrease  in  number  since  1830  is  admitted  by  every  one,  in- 
cluding the  natives  themselves,  and  is  still  steadily  progressing. 
Although  it  has  hitherto  been  found  impossible  to  take  an  actual 
census  of  the  natives,  their  numbers  were  carefully  estimated  by 
residents  in  many  districts.  The  result  seems  trustworthy,  and 
shows  that  during  the  fourteen  years,  previous  to  1858,  the  de- 
crease was  19.42  per  cent.  Some  of  the  tribes,  thus  carefully 
examined,  lived  above  a  hundred  miles  apart,  some  on  the  coast, 
some  inland;  and  their  means  of  subsistence  and  habits  differed 
to  a  certain  extent  (p.  28).  The  total  number  in  1858  was  be- 
lieved to  be  53,700,  and  in  1872,  after  a  second  interval  of  fourteen 
years,  another  census  was  taken,  and  the  number  is  given  as 
only  36,359,  showing  a  decrease  of  32.29  per  cent.!^  Mr.  Fenton, 
after  showing  in  detail  the  insufficiency  of  the  various  causes, 
usually  assigned  in  explanation  of  this  extraordinary  decrease, 
such  as  new  diseases,  the  profligacy  of  the  women,  drunkenness, 
wars,  &c.,  concludes  on  weighty  grounds  that  it  depends  chiefly 
on  the  unproductiveness  of  the  women,  and  on  the  extraordinary 
mortality  of  the  young  children  (pp.  31,  34).  In  proof  of  this  he 
shows  (p.  33)  that  in  1844  there  was  one  non-adult  for  every 
2.57  adults;  whereas  in  1858  there  was  only  one  non-adult  for 
every  3.27  adults.  The  mortality  of  the  adults  is  also  great.  He 
adduces  as  a  further  cause  of  the  decrease  the  inequality  of  the 
sexes;  for  fewer  females  are  born  than  males.  To  this  latter 
point,  depending  perhaps  on  a  widely  distinct  cause,  I  shall  re- 
turn in  a  future  chapter.  Mr.  Fenton  contrasts  with  astonish- 
ment the  decrease  in  New  Zealand  with  the  increase  in  Ireland; 


39  For  these  cases,  see  Bonwick's  'Daily  Life  of  the  Tasmanians,' 
1870,  p.  90  and  the  'Last  of  the  Tasmanians,'  1870,  p.  386. 

*o  'Observations  on  the  Aboriginal  Inhabitants  of  New  Zealand,'  pub- 
lished by  the  Government,  1859. 

*i  'New  Zealand,'  by  Alex.  Kennedy,  1873,  p.  47. 


THE    EXTINCTION    OF    RACES. 


countries  not  very  dissimilar  in  climate,  and  where  the  inhabi- 
tants now  follow  nearly  similar  habits.  The  Maories  themselves 
(p.  85)  "attribute  their  decadence,  in  some  measure,  to  the  intro- 
"duction  of  new  food  and  clothing,  and  the  attendant  change  of 
"habits;"  and  it  will  be  seen,  when  we  consider  the  influence  of 
changed  conditions  on  fertility,  that  they  are  probably  right. 
The  diminution  began  between  the  years  1830  and  1840;  and  Mr. 
Fenton  shows  (p.  40)  that  about  1830,  the  art  of  manufacturing 
putrid  corn  (maize),  by  long  steeping  in  water,  was  discovered 
and  largely  practiced;  and  this  proves  that  a  change  of  habits  was 
beginning  amongst  the  natives,  even  when  New  Zealand  was  only 
thinly  inhabited  by  Europeans.  When  I  visited  the  Bay  of  Is- 
lands in  1835,  the  dress  and  food  of  the  inhabitants  had  already 
been  much  modified:  they  raised  potatoes,  maize,  and  other  agri- 
cultural produce,  and  exchanged  them  for  English  manufactured 
goods  and  tobacco. 

It  is  evident  from  many  statements  in  the  life  of  Bishop  Patte- 
son,*^  that  the  Melanesians  of  the  New  Hebrides  and  neighboring 
archipelagoes,  suffered  to  an  extraordinary  degree  in  health,  and 
perished  in  large  numbers,  when  they  were  removed  to  New 
Zealand,  Norfolk  Island,  and  other  salubrious  places,  in  order  to 
be  educated  as  missionaries. 

The  decrease  of  the  native  population  of  the  Sandwich  Islands 
is  as  notorious  as  that  of  New  Zealand.  It  has  been  roughly 
estimated  by  those  best  capable  of  judging,  that  when  Cook  dis- 
covered the  Islands  in  1779,  the  population  amounted  to  about 
300,000.  According  to  a  loose  census  in  1823,  the  numbers  then 
were  142,050.  In  1832,  and  at  several  subsequent  periods,  an  ac- 
curate census  was  officially  taken,  but  I  have  been  able  to  obtain 
only  the  following  returns: 


Year. 


Native    Population. 

(Except  during  1832 
and  1836,  when  the 
few  foreigners  in 
the  islands  were 
included). 


Annual  rate  of  de- 
crease per  cent., 
assuming  it  to 
have  been  uniform 
between  the  suc- 
cessive censuses ; 
these  censuses  be- 
ing taken  at  irreg- 
ular   intervals. 


1832 
1836 
1853 
1860 
1866 
1872 


130.313 


108.579  i 

71,019  [ 

67,084  [ 

58,765  [ 

51,531  r 


4.46 
2.47 
0.81 
2.18 

i.n 


*2  'Life  of  J.  C.  Patteson,'   by  C.  M.  Younge,  1874;  see  more  especially 
vol.  i.  p.   530. 
13 


182  THE  DESCENT  OF  MAN. 

We  here  see  that  in  the  interval  of  forty  years,  between  1832 
and  1872,  the  population  has  decreased  no  less  than  sixty-eight 
per  cent.!  This  has  been  attributed  by  most  writers  to  the 
profligacy  of  the  women,  to  former  bloody  wars,  and  to  the  severe 
labor  imposed  on  conquered  tribes  and  to  newly  introduced  dis- 
eases, which  have  been  on  several  occasions  extremely  destruc- 
tive. No  doubt  these  and  other  such  causes  have  been  highly  ef- 
ficient, and  may  account  for  the  extraordinary  rate  of  decrease 
between  the  years  1832  and  1836;  but  the  most  potent  of  all  the 
causes  seems  to  be  lessened  fertility.  According  to  Dr.  Ruschen- 
berger  of  the  U.  S.  Navy,  who  visited  these  islands  between  1835 
and  1837,  in  one  district  of  Hawaii,  only  twenty-five  men  out  of 
1134,  and  in  another  district  only  ten  out  of  637,  had  a  family 
with  as  many  as  three  children.  Of  eighty  married  women,  only 
thirty-nine  had  ever  borne  children;  and  "the  official  report  gives 
"an  average  of  half  a  child  to  each  married  couple  in  the  whole 
"island."  This  is  almost  exactly  the  same  average  as  with  the 
Tasmanians  at  Oyster  Cove.  Jarves,  who  published  his  History 
in  1843,  says  that  "families  who  have  three  children  are  freed 
"from  all  taxes;  those  having  more,  are  rewarded  by  gifts  of  land 
"and  other  encouragements."  This  unparalleled  enactment  by 
the  government  well  shows  how  infertile  the  race  had  become. 
The  Rev.  A.  Bishop  stated  in  the  Hawaiian  'Spectator'  in  1839,  that 
a  large  proportion  of  the  children  die  at  early  ages,  and  Bishop 
Staley  informs  me  that  this  is  still  the  case,  just  as  in  New 
Zealand.  This  has  been  attributed  to  the  neglect  of  the  children 
by  the  women,  but  it  is  probably  in  large  part  due  to  innate 
weakness  of  constitution  in  the  children,  in  relation  to  the  les- 
sened fertility  of  their  parents.  There  is,  moreover,  a  further  re- 
semblance to  the  case  of  New  Zealand,  in  the  fact  that  there  is  a 
large  excess  of  male  over  female  births:  the  census  of  1872  gives 
31,650  males  to  25,247  females  of  all  ages,  that  is  125.36  males  for 
every  100  females;  whereas  in  all  civilized  countries  the  females 
exceed  the  males.  No  doubt  the  profligacy  of  the  women  may  in 
part  account  for  their  small  fertility;  but  their  changed  habits  of 
life  is  a  much  more  probable  cause,  and  which  will  at  the  same 
time  account  for  the  increased  mortality,  especially  of  the  chil- 
dren. The  islands  were  visited  by  Cook  in  1779,  by  Vancouver  in 
1794,  and  often  subsequently  by  whalers.  In  1819  missionaries 
arrived,  and  found  that  idolatry  had  been  already  abolished,  and 
other  changes  effected  by  the  king.  After  this  period  there  was 
a  rapid  change  in  almost  all  the  habits  of  life  of  the  natives,  and 
they  soon  became  "the  most  civilized  of  the  Paciflc  Islanders." 
One  of  my  informants,  Mr.  Coan,  who  was  born  on  the  islands, 
remarks  that  the  natives  have  undergone  a  greater  change  in 
their  habits  of  life  in  the  course  of  fifty  years  than  Englishmen 
during    a    thousand    years.    From    information    received    from 


THE    EXTINCTION    OF    RACES.  ISS 

Bishop  Staley,  it  does  not  appear  that  the  poorer  classes  have 
ever  much  changed  their  diet,  although  many  new  kinds  of  fruit 
have  been  introduced,  and  the  sugar-cane  is  in  universal  use. 
Owing,  however,  to  their  passion  for  imitating  Europeans,  they 
altered  their  manner  of  dressing  at  an  early  period,  and  the  use 
of  alcoholic  drinks  became  very  general.  Although  these  changes 
appear  inconsiderable,  I  can  well  believe,  from  what  is  known 
with  respect  to  animals,  that  they  might  suflBce  to  lessen  the  fer- 
tility of  the  natives.^ 

Lastly,  Mr,  Macnamara  states^  that  the  low  and  degraded  in- 
habitants of  the  Andaman  Islands,  on  the  eastern  side  of  the  Gulf 
of  Bengal,  are  "eminently  susceptible  to  any  change  of  climate: 
"in  fact,  take  them  away  from  their  island  homes,  and  they  are 
"almost  certain  to  die,  and  that  independently  of  diet  or  ex- 
"traneous  influences."  He  further  states  that  the  inhabitants  of 
the  valley  of  Nepal,  which  is  extremely  hot  in  summer,  and  also 
the  various  hill-tribes  of  India,  suffer  from  dysentery  and  fever 
when  on  the  plains;  and  they  die  if  they  attempt  to  pass  the 
whole  year  there. 

We  thus  see  that  many  of  the  wilder  races  of  man  are  apt  to 
suffer  much  in  health  when  subjected  to  changed  conditions 
or  habits  of  life,  and  not  exclusively  from  being  transported  to 
a  new  climate.  Mere  alterations  in  habits,  which  do  not  appear 
injurious  in  themselves,  seem  to  have  this  same  effect;  and  in 
several  cases  the  children  are  particularly  liable  to  suffer.  It 
has  often  been  said,  as  Mr.  Macnamara  remarks,  that  man  can 
resist  with  impunity  the  greatest  diversities  of  climate  and  other 
changes;  but  this  is  true  only  of  the  civilized  races.  Man  in 
his  wild  condition  seems  to  be  in  this  respect  almost  as  sus- 
ceptible as  his  nearest  allies,  the  anthropoid  apes,  which  have 
never  yet  survived  long,  when  removed  from  their  native 
country. 

Lessened  fertility  from  changed  conditions,  as  in  the  case  of 
the  Tasmanians,  Maories,  Sandwich  Islanders,  and  apparently 
the  Australians,  is  still  more  interesting  than  their  liability  to 
ill-health  and  death;  for  even  a  slight  degree  of  infertility  com- 

*3  The  foregoing  statements  are  taken  chiefly  from  the  following 
works:  'Jarves'  History  of  the  Hawaiian  Islands,'  1843,  p.  400-407. 
Cheever,  'Life  in  the  Sandwich  Islands,'  1851,  p.  277.  Ruschenberger  is 
quoted  by  Bonwick,  'Last  of  the  Tasmanians,'  1870,  p.  378.  Bishop  is 
quoted  by  Sir  E.  Belcher,  'Voyage  Round  the  World,'  1843,  vol.  i.,  p. 
272.  I  owe  the  census  of  the  several  years  to  the  kindness  of  Mr.  Coan, 
at  the  request  of  Dr.  Youmans  of  New  York,  and  in  most  ca&es  I  have 
compared  the  Youmans  figures  with  those  given  in  several  of  the 
above-named  works.  I  have  omitted  the  census  for  1850,  as  I  have 
seen  two  widely  different  numbers  given. 

^  'The  Indian  Medical  Gazette,'  Nov.  1,  1871,  p.  240. 


Ig4  THE  DESCENT  OP  MAN. 

"bined  with  those  other  causes  which  tend  to  checl^  the  increase 
of  every  population,  would  sooner  or  later  lead  to  extinction. 
The  diminution  of  fertility  may  be  explained  in  some  cases  by 
the  profligacy  of  the  women  (as  until  lately  with  the  Tahitians) 
but  Mr.  Penton  has  shown  that  this  explanation  by  no  means  suf- 
fices with  the  New  Zealanders,  nor  does  it  with  the  Tasmanians. 

In  the  paper  above  quoted,  Mr.  Macnamara  gives  reasons  for 
believing  that  the  inhabitants  of  districts  subject  to  malaria  are 
apt  to  be  sterile;  but  this  cannot  apply  in  several  of  the  above 
cases.  Some  writers  have  suggested  that  the  aborigines  of  is- 
lands have  suffered  in  fertility  and  health  from  long  continued 
inter-breeding;  but  in  the  above  cases  infertility  has  coincided 
too  closely  with  the  arrival  of  Europeans  for  us  to  admit  this 
explanation.  Nor  have  we  at  present  any  reason  to  believe 
that  man  is  highly  sensitive  to  the  evil  effects  of  inter-breeding, 
especially  in  areas  so  large  as  New  Zealand,  and  the  Sandwich 
archipelago  with  its  diversified  stations.  On  the  contrary,  it  is 
known  that  the  present  inhabitants  of  Norfolk  Island  are  nearly 
all  cousins  or  near  relations,  as  are  the  Todas  in  India,  and  the 
inhabitants  of  some  of  the  Western  Islands  of  Scotland;  and  yet 
they  seem  not  to  have  suffered  in  fertility.^ 

A  much  more  probable  view  is  suggested  by  the  analogy  of  the 
lower  animals.  The  reproductive  system  can  be  shown  to  be  sus- 
ceptible to  an  extraordinary  degree  (though  why  we  know  not) 
to  changed  conditions  of  life;  and  this  susceptibility  leads  both 
to  beneficial  and  to  evil  results.  A  large  collection  of  facts  on 
this  subject  is  given  in  chap,  xviii.  of  vol.  ii.  of  my  'Variation  of 
Animals  and  Plants  under  Domestication,'  I  can  here  give  only 
the  briefest  abstract;  and  every  one  interested  in  the  subject  may 
consult  the  above  work.  Very  slight  changes  increase  the  health, 
vigor  and  fertility  of  most  or  all  organic  beings,  whilst  other 
changes  are  known  to  render  a  large  number  of  animals  sterile. 
One  of  the  most  familiar  cases,  is  that  of  tamed  elephants,  not 
breeding  in  India;  though  they  often  breed  in  Ava,  where  the 
females  are  allowed  to  roam  about  the  forests  to  some  extent,  and 
are  thus  placed  under  more  natural  conditions.  The  case  of 
various  American  monkeys,  both  sexes  of  which  have  been  kept 
for  many  yekrs  together  in  their  own  countries,  and  yet  have 
very  rarely  or  never  bred,  is  a  more  apposite  instance,  because  of 
their  relationship  to  man.  It  is  remarkable  how  slight  a  change 
in  the  conditions  often  induces  sterility  in  a  wil'd  animal  when 
eaptured;    and  this  is  the  more  strange  as  all  our  domesticated 

*^  On  the  close  relationship  of  the  Norfolk  Islanders,  see  Sir  W. 
Denison,  'Varieties  of  Vice-Regal  Life,'  vol.  i.  1870,  p.  410.  For  the 
Todas,  see  Col.  Marshall's  work,  1873,  p.  110.  For  the  Western  Islands 
of  Scotland,  Dr.  Mitchell,  'Edinburgh  Medical  Journal,'  March  to 
June,   1865. 


THE    EXTINCTION   OF   RACES.  185 

animals  have  become  more  fertile  than  they  were  in  a  state  of 
nature;  and  some  of  them  can  resist  the  most  unnatural  condi- 
tions with  undiminished  fertility.*^  Certain  groups  of  animals 
are  much  more  liable  than  others  to  be  affected  by  captivity; 
and  generally  all  the  species  of  the  same  group  are  affected  in 
the  same  manner.  But  sometimes  a  single  species  in  a  group 
is  rendered  sterile,  whilst  the  others  are  not  so;  on  the  other 
hand,  a  single  species  may  retain  its  fertility  whilst  most  of  the 
others  fail  to  breed.  The  males  and  females  of  some  species 
when  confined,  or  when  allowed  to  live  almost,  but  not  quite  free, 
in  their  native  country,  never  unite;  others  thus  circumstanced 
frequently  unite  but  never  produce  offspring;  others  again  pro- 
duce some  offspring,  but  fewer  than  in  a  state  of  nature;  and 
as  bearing  on  the  above  cases  of  man,  it  is  important  to  remark 
that  the  young  are  apt  to  be  weak  and  sickly,  or  malformed,  and 
to  perish  at  an  early  age. 

Seeing  how  general  is  this  law  of  the  susceptibility  of  the 
reproductive  system  to  changed  conditions  of  life,  and  that  it 
holds  good  with  our  nearest  allies,  the  Quadrumana,  I  can  hardly 
doubt  that  it  applies  to  man  in  his  primeval  state.  Hence  if 
savages  of  any  race  are  induced  suddenly  to  change  their  habits 
of  life,  they  become  more  or  less  sterile,  and  their  young  off- 
spring suffer  in  health,  in  the  same  manner  and  from  the  same 
cause,  as  do  the  elephant  and  hunting-leopard  in  India,  many 
monkeys  in  America,  and  a  host  of  animals  of  all  kinds,  on  re- 
moval from  their  natural  conditions. 

We  can  see  why  it  is  that  aborigines,  who  have  long  inhabited 
islands,  and  who  must  have  been  long  exposed  to  nearly  uniform 
conditions,  should  be  specially  affected  by  any  change  in  their 
habits,  as  seems  to  be  the  case.  Civilized  races  can  certainly 
resist  changes  of  all  kinds  far  better  than  savages;  and  in  this 
respect  they  resemble  domesticated  animals,  for  though  the  lat- 
ter sometimes  suffer  in  health  (for  instance  European  dogs  in 
India),  yet  they  are  rarely  rendered  sterile,  though  a  few  such 
instances  have  been  recorded.*^  The  immunity  of  civilized  races 
and  domesticated  animals  is  probably  due  to  their  having  been 
subjected  to  a  greater  extent,  and  therefore  having  grown  some- 
what more  accustomed,  to  diversified  or  varying  conditions,  than 
the  majority  of  wild  animals;  and  to  their  having  formerly 
immigrated  or  been  carried  from  country  to  country,  and  to  dif- 
ferent families  or  sub-races  having  inter-crossed.  It  appears 
that  a  cross  with  civilized  races  at  once  gives  to  an  aboriginal 
race  an  immunity  from  the  evil  consequences  of  changed  condi- 


^  For  the   evidence   on  this   head,   see   'Variation   of  Animals,'  &c., 
vol.  ii.  p.  111. 
*^  'Variation  of  Animals,'  &c.,  vol.  ii.,  p.  16. 


186  THE  DESCENT  OP  MAN. 

tions.  Tlius  tlie  crossed  offspring  from  the  Tahitians  and  Eng- 
lish, when  settled  in  Pitcairn  Island,  increased  so  rapidly  that 
the  island  was  soon  overstocked;  and  in  June  1856  they  were  re- 
moved to  Norfolk  Island.  They  then  consisted  of  60  married 
persons  and  134  children,  making  a  total  of  194.  Here  they  like- 
wise increased  so  rapidly,  that  although  sixteen  of  them  returned 
to  Pitcairn  Island  in  1859,  they  numbered  in  January  1868,  300 
souls,  the  males  and  females  being  in  exactly  equal  numbers. 
What  a  contrast  does  this  case  present  with  that  of  the  Tas- 
manians;  the  Norfolk  Islanders  increased  in  only  twelve  and  a 
half  years  from  194  to  300;  whereas  the  Tasmanians  decreased 
during  fifteen  years  from  120  to  46,  of  which  latter  number  only 
ten  were  children.*^ 

So  again  in  the  interval  between  the  census  of  1866  and  1872 
the  natives  of  full  blood  in  the  Sandwich  Islands  decreased  by 
8081,  whilst  the  half-castes,  who  are  believed  to  be  healthier,  in- 
creased by  847;  but  I  do  not  know  whether  the  latter  number 
includes  the  offspring  from  the  half-castes,  or  only  the  half- 
castes  of  the  first  generation. 

The  cases  which  I  have  here  given  all  relate  to  aborigines, 
who  have  been  subjected  to  new  conditions  as  the  result  of 
the  immigration  of  civilized  men.  But  sterility  and  ill-health 
would  probably  follow,  if  savages  were  compelled  by  any  cause, 
such  as  the  inroad  of  a  conquering  tribe,  to  desert  their  homes 
and  to  change  their  habits.  It  is  an  interesting  circumstance 
that  the  chief  check  to  wild  animals  becoming  domesticated, 
which  implies  the  power  of  their  breeding  freely  when  first  cap- 
tured, and  one  chief  check  to  wild  men,  when  brought  into  con- 
tact with  civilization,  surviving  to  form  a  civilized  race,  is  the 
same,  namely,  sterility  from  changed  conditions  of  life. 

Finally,  although  the  gradual  decrease  and  ultimate  extinc- 
tion of  the  races  of  man  is  a  highly  complex  problem,  depending 
on  many  causes  which  differ  in  different  places  and  at  different 
times;  it  is  the  same  problem  as  that  presented  by  the  extinc- 
tion of  one  of  the  higher  animals — of  the  fossil  horse,  for  in- 
stance, which  disappeared  from  South  America,  soon  afterwards 
to  be  replaced,  within  the  same  districts,  by  countless  troops 
of  the  Spanish  horse.  The  New  Zealander  seems  conscious  of 
this  parallelism,  for  he  compares  his  future  fate  with  that  of 
the  native  rat  now  almost  exterminated  by  the  European  rat. 
Though  the  difficulty  is  great  to  our  imagination,  and  really 
great,  if  we  wish  to  ascertain  the  precise  causes  and  their  man- 

*8  These  details  are  taken  from  'The  Mutineers  of  the  "Bounty,"  ' 
by  Lady  Belcher,  1870;  and  from  'Pitcairn  Island,'  ordered  to  be 
printed  by  the  House  of  Commons,  May  29th,  1863.  The  following- 
statements  about  the  Sandwich  Islanders  are  from  the  'Honolulu  Ga- 
zette,' and  from  M>.  Coan. 


THE  FORMATION  OF  RACES.  187 

ner  of  action,  it  ought  not  to  be  so  to  our  reason,  as  long  as  we 
keep  steadily  in  mind  that  the  increase  of  each  species  and  each 
race  is  constantly  checked  in  various  ways;  so  that  if  any  new 
check,  even  a  slight  one,  be  superadded,  the  race  will  surely  de- 
crease in  number;  and  decreasing  numbers  will  sooner  or  later 
lead  to  extinction;  the  end,  in  most  cases,  being  promptly  deter- 
mined by  the  inroads  of 'conquering  tribes. 

On  the  Formation  of  the  Races  of  Man. — In  some  cases  the 
crossing  of  distinct  races  has  led  to  the  formation  of  a  new  race. 
The  singular  fact  that  Europeans  and  Hindoos,  who  belong  to 
the  same  Aryan  stock,  and  speak  a  language  fundamentally  the 
same,  differ  widely  in  appearance,  whilst  Europeans  differ  but 
little  from  Jews,  who  belong  to  the  Semitic  stock,  and  speak 
quite  another  language,  has  been  accounted  for  by  Broca,*^ 
through  certain  Aryan  branches  having  been  largely  crossed  by 
indigenous  tribes  during  their  wide  diffusion.  When  two  races 
in  close  contact  cross,  the  first  result  is  a  heterogeneous  mixture: 
thus  Mr.  Hunter,  in  describing  the  Santali  or  hill-tribes  of  India, 
says  that  hundreds  of  imperceptible  gradations  may  be  traced 
"from  the  black,  squat  tribes  of  the  mountains  to  the  tall  olive- 
"colored  Brahman,  with  his  intellectual  brow,  calm  eyes,  and 
"high  but  narrow  head;"  so  that  it  is  necessary  in  courts  of 
justice  to  ask  the  witnesses  whether  they  are  Santalis  or  Hin- 
doos.^" Whether  a  heterogeneous  people,  such  as  the  inhabit- 
ants of  some  of  the  Polynesian  Islands,  formed  by  the  crossing 
of  two  distinct  races,  with  few  or  no  pure  members  left,  would 
ever  become  homogeneous,  is  not  known  from  direct  evidence. 
But  as  with  our  domesticated  animals,  a  cross-breed  can  cer- 
tainly be  fixed  and  made  uniform  by  careful  selection^^  in  the 
course  of  a  few  generations,  we  may  infer  that  the  free  inter- 
crossing of  a  heterogeneous  mixture  during  a  long  descent  would 
supply  the  place  of  selection,  and  overcome  any  tendency  to 
reversion;  so  that  the  crossed  race  would  ultimately  become 
homogeneous,  though  it  might  not  partake  in  an  equal  degree 
of  the  characters  of  the  two  parent-races. 

Of  all  the  differences  between  the  races  of  man,  the  color  of 
the  skin  is  the  most  conspicuous  and  one  of  the  best  marked.  It 
was  formerly  thought  that  differences  of  this  kind  could  be 
accounted  for  by  long  exposure  to  different  climates;  but  Pal- 
las first  showed  that  this  is  not  tenable,  and  he  has  since  been 


^  'On  Anthropology,'  translation  'Anthropolog-.  Review,'  Jan.  1868, 
p.  38. 

5"  'The  Annals  of  Rural  Bengal,'  1868  p.  134. 

^^  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
p.  95. 


188  THE  DESCENT  OF  MAN. 

followed  by  almost  all  anthropologists.^^  This  view  has  been 
rejected  chiefly  because  the  distribution  of  the  variously  col- 
ored races,  most  of  whom  must  have  long  inhabited  their  present 
homes,  does  not  coincide  with  corresponding  differences  of  cli- 
mate. Some  little  weight  may  be  given  to  such  cases  as  that  of 
the  Dutch  families,  who,  as  we  hear  on  excellent  authority.^" 
have  not  undergone  the  least  change  of  color  after  residing  for 
three  centuries  in  South  Africa.  An  argument  on  the  same  side 
may  likewise  be  drawn  from  the  uniform  appearance  in  various 
parts  of  the  world  of  gipsies  and  Jews,  though  the  uniformity  of 
the  latter  has  been  somewhat  exaggerated.^*  A  very  damp  or  a 
very  dry  atmosphere  has  been  supposed  to  be  more  influential  in 
modifying  the  color  of  the  skin  than  mere  heat;  but  as  D'Or- 
bigny  in  South  America,  and  Livingstone  in  Africa,  arrived  at 
diametrically  opposite  conclusions  with  respect  to  dampness  and 
dryness,  any  conclusion  on  this  head  must  be  considered  as  very 
doubtful.'" 

Various  facts,  which  I  have  given  elsewhere,  prove  that  the 
color  of  the  skin  and  hair  is  sometimes  correlated  in  a  surpris- 
ing manner  with  a  complete  immunity  from  the  action  of  certain 
vegetable  poisons,  and  from  the  attacks  of  certain  parasites. 
Hence  it  occurred  to  me,  that  negroes  and  other  dark  races 
might  have  acquired  their  dark  tints  by  the  darker  Individuals 
escaping  from  the  deadly  influence  of  the  miasma  of  their  native 
countries,  during  a  long  series  of  generations. 

I  afterwards  found  that  this  same  idea  had  long  ago  occurred 
to  Dr.  Wells.'°  It  has  long  been  known  that  negroes,  and  even 
mulattoes,  are  almost  completely  exempt  from  the  yellow  fever, 
so  destructive  in  tropical  America."  They  likewise  escape  to  a 
large  extent  the  fatal  intermittent  fevers,  that  prevail  along  at 
least  2600  miles  of  the  shores  of  Africa,  and  which  annually 
cause  one-fifth  of  the  white  settlers  to  die,  and  another  fifth  to 


52  Pallas,  'Act.  Acad.  St.  Petersburg,'  1780,  part  ii.  p.  69.  He  was  fol- 
lowed by  Hudolphi,  in  his  'Beytrage  zur  Anthropologie,'  1812.  An  ex- 
cellent summary  of  the  evidence  is  given  by  Godron,  'De  I'Espece,' 
1859,  vol.  ii.  p.  246,  &c. 

53  Sir  Andrew  Smith,  as  quoted  by  Knox,  'Races  of  Man,'  1850,  p.  473. 
^*  See  De  Quatrefages  on  this  head,  'Revue  des  Cours  Scientifiques,' 

Oct.  17,  1868,  p.  731. 

^5  Livingstone's  'Travels  and  Researches  in  S.  Africa,'  1857,  pp.  338, 
329.    D'Orbigny,   as  quoted  by  Godron,   'De  I'Espece,'  vol.  ii.  p.  266. 

58  See  a  paper  read  before  the  Royal  Soc.  in  1813,  and  published  in 
his  Essays  in  1818.  I  have  given  an  account  of  Dr.  Wells'  views  in 
the  Historical  Sketch  (p.  xvi.)  to  my  'Origin  of  Species.'  Various 
cases  of  color  correlated  with  constitutional  peculiarities  are  given  in 
my  'Variation  of  Animals  under  Domestication,'  vol.   ii.  pp.  227,   335. 

57  See,  for  instance,  Nott  and  Gliddon,  'Types  of  Mankind,'  p.  68. 


THE    FORMATION    OF    RACES.  189 

return  liome  inralided/^  This  immunity  in  the  negro  seems  to 
be  partly  inherent,  depending  on  some  unknown  peculiarity  of 
constitution,  and  partly  the  result  of  acclimatization.  Pouchet'* 
states  that  the  negro  regiments  recruited  near  the  Soudan,  and 
borrowed  from  the  Viceroy  of  Egypt  for  the  Mexican  war, 
escaped  the  yellow-fever  almost  equally  v/ith  the  negroes  origin- 
ally brought  from  various  parts  of  Africa  and  accustomed  to  the 
climate  of  the  "West  Indies.  That  acclimatization  plays  a  part, 
is  shown  by  the  many  cases  in  which  negroes  have  become  some- 
what liable  to  tropical  fevers,  after  having  resided  for  some  time 
in  a  colder  climate.'''^  The  nature  of  the  climate  under  which  the 
white  races  have  long  resided,  likewise  has  some  influence  on 
them;  for  during  the  fearful  epidemic  of  yellow-fever  in  Dem- 
erara  during  1837,  Dr.  Blair  found  that  the  death-rate  of  the 
immigrants  was  proportional  to  the  latitude  of  the  country 
whence  they  had  come.  With  the  negro  the  immunity,  as  far  as 
it  is  the  result  of  acclimatization,  implies  exposure  during  a 
prodigious  length  of  time;  for  the  aborigines  of  tropical  America 
who  have  resided  there  from  time  immemorial,  are  not  exempt 
from  yellow  fever;  and  the  Rev.  H.  B.  Tristram  states,  that  there 
are  districts  in  Northern  Africa  which  the  native  inhabitants  are 
compelled  annually  to  leave,  though  the  negroes  can  remain  with 
safety. 

That  the  immunity  of  the  negro  is  in  any  degree  correlated 
with  the  color  of  his  skin  is  a  mere  conjecture:  it  may  be  cor- 
related with  some  difference  in  his  blood,  nervous  system,  or 
other  tissues.  Nevertheless,  from  the  facts  above  alluded  to,  and 
from  some  connection  apparently  existing  between  complexion 
and  a  tendency  to  consumption,  the  conjecture  seemed  to  me  not 
improbable.     Consequently  I  endeavored,  with  but  little  success,®^ 

58  Major  Tulloch,  in  a  paper  read  before  the  Statistical  Society, 
April  20th,  1840,  and  given  in  the  'Athenaeum,'  1840,  p.  353. 

59  'The  Plurahty  of  the  Human  Race,'   (translat.),  1864,  p.   60. 

60  Quatrefages,  'Unite  de  I'Espece  Humaine,'  1861,  p.  205.  Waltz, 
'Introduct.  to  Anthropology,'  translat.  vol.  i.  1883,  p.  124.  Livingstone 
gives  analogous  cases  in  his  'Travels.' 

51  In  the  spring  of  1862  I  obtained  permission  from  the  Director-Gen- 
eral of  the  Medical  department  of  the  Army,  to  transmit  to  the  sur- 
geons of  the  various  regiments  on  foreign  service  a  blank  table,  with 
the  following  appended  remarks,  but  I  have  received  no  returns.  "As 
"several  well-marked  cases  have  been  recorded  with  our  domestic 
"animals  of  a  relation  between  the  color  of  the  dermal  appendages 
"and  the  constitution;  and  it  being  notorious  that  there  is  some  lim- 
"ited  degree  of  relation  between  the  color  of  the  races  of  man  and 
"the  climate  inhabited  by  them;  the  following  investigation  seems 
"worth  consideration.  Namely,  whether  there  is  any  relation  in  Eu- 
"ropeans  between  the  color  of  their  hair,  and  their  liability  to  the  dis- 
"eases    of    tropical   countries.    If   the    surgeons    of    the   several   regi- 


190  THE  DESCENT  OF  MAN. 

to  ascertain  how  far  it  holds  good.  The  late  Dr.  Daniell,  who 
had  long  lived  on  the  West  Coast  of  Africa,  told  me  that  he  did 
not  believe  in  any  such  relation.  He  was  himself  unusually  fair, 
and  had  withstood  the  climate  in  a  wonderful  manner.  When 
he  first  arrived  as  a  boy  on  the  coast,  an  old  and  experienced 
negro  chief  predicted  from  his  appearance  that  this  would  prove 
the  case.  Dr.  Nicholson,  of  Antigua,  after  having  attended  to  this 
subject,  writes  to  me  that  he  does  not  think  that  dark-colored  Eu- 
ropeans escape  the  yellow-fever  more  than  those  that  are  light- 
colored.  Mr.  J.  M.  Harris  altogether  denies  that  Europeans  with 
dark  hair  withstand  a  hot  climate  better  than  other  men:  on  the 
contrary,  experience  has  taught  him  in  making  a  selection  of 
men  for  service  on  the  coast  of  Africa,  to  choose  those  with  red 
hair."-  As  far,  therefore,  as  these  slight  indications  go,  there 
seems  no  foundation  for  the  hypothesis,  that  blackness  has  re- 
sulted from  the  darker  and  darker  individuals  having  survived 
better  during  long  exposure  to  fever-generating  miasma. 

Dr.  Sharpe  remarks,^  that  a  tropical  sun,  which  burns  and 
blisters  a  white  skin,  does  not  injure  a  black  one  at  all;  and,  as 
he  adds,  this  is  not  due  to  habit  in  the  individual,  for  children 
only  six  or  eight  months  old  are  often  carried  about  naked,  and 
are  not  affected.    I  have  been  assured  by  a  medical  man,  that 

"ments,  when  stationed  in  unhealthy  tropical  districts,  would  be  so 
"good  as  first  to  count,  as  a  standard  of  comparison,  how  many  men, 
"in  the  force  whence  the  sick  are  drawn,  have  dark  and  light- 
"colored  hair,  and  hair  of  intermediate  or  doubtful  tints;  and  if  a 
"similar  account  were  kept  by  the  same  medical  gentlemen,  of  all 
"the  men  who  suffered  from  malarious  and  yellow  fevers,  or  from 
"dysentery,  it  would  soon  be  apparent,  after  some  thousand  cases  had 
"been  tabulated,  whether  there  exists  any  relation  between  the  color 
"of  the  hair  and  constitutional  liability  to  tropical  diseases.  Per- 
"haps  no  such  relation  would  be  discovered,  but  the  investigation  is 
"well  worth  making.  In  case  any  positive  result  were  obtained,  it 
"might  be  of  some  practical  use  in  selecting  men  for  any  particular 
"service.  Theoretically  the  result  would  be  of  high  interest,  as  indl- 
"cating  one  means  by  which  a  race  of  men  inhabiting  from  a  remote 
"period  an  unhealthy  tropical  climate,  might  have  become  dark-col- 
"ored  by  the  better  preservation  of  dark-haired  or  dark-complexioned 
"individuals   during   a  long   succession   of  generations." 

"2  'Anthropological  Review,'  Jan.  1866,  p.  xxi.  Dr.  Sharpe  also  says 
with  respect  to  India  ('Man  a  Special  Creation,'  1873,  p.  118),  that  "it 
"has  been  noticed  by  some  medical  officers  that  Europeans  with  light 
"hair  and  florid  complexions  suffer  less  from  diseases  of  tropical 
"countries  than  persons  with  dark  hair  and  sallow  complexions; 
"and,  so  far  as  I  know,  there  appear  to  be  good  grounds  for  this 
"remark."  On  the  other  hand,  Mr.  Heddle,  of  Sierra  Leone,  "who 
"has  had  more  clerks  killed  under  him  than  any  other  man,"  by  the 
climate  of  the  West  African  Coast  (W.  Reade,  'African  Sketch  Book,' 
vol.  ii.  p.  522),  holds  a  directly  opposite  view,  as  does  Capt.  Burton. 

«3  'Man  a  Special  Creation,'  1873,  p.  119. 


THE    FORMATION    OF    RACES.  191 

some  years  ago  during  each  summer,  but  not  during  the  winter, 
his  hands  became  marked  with  light  brown  patches,  like,  al- 
though larger  that  freckles,  and  that  these  patches  were  never 
affected  by  sun-burning,  whilst  the  white  parts  of  his  skin  have 
on  several  occasions  been  much  inflamed  and  blistered.  With 
the  lower  animals  there  is,  also,  a  constitutional  difference  in  lia- 
bility to  the  action  of  the  sun  between  those  parts  of  the  skin 
clothed  with  white  hair  and  other  parts.°^  Whether  the  saving 
of  the  skin  from  being  thus  burnt  is  of  sufficient  importance  to 
account  for  a  dark  tint  having  been  gradually  acquired  by  man 
through  natural  selection,  I  am  unable  to  judge.  If  it  be  so,  we 
should  have  to  assume  that  the  natives  of  tropical  America  have 
lived  there  for  a  much  shorter  time  than  the  negroes  in  Africa, 
or  the  Papuans  in  the  southern  parts  of  the  Malay  archipelago, 
just  as  the  lighter-colored  Hindoos  have  resided  in  India  for  a 
shorter  time  than  the  darker  aborigines  of  the  central  and  south- 
ern parts  of  the  peninsula. 

Although  with  our  present  knowledge  we  cannot  account  for 
the  differences  of  color  in  the  races  of  man,  through  any  advan- 
tage thus  gained,  or  from  the  direct  action  of  climate;  yet  we 
must  not  quite  ignore  the  latter  agency,  for  there  is  good  reason 
to  believe  that  some  inherited  effect  is  thus  produced.^^ 

We  have  seen  in  the  second  chapter  that  the  conditions  of  life 
affect  the  development  of  the  bodily  frame  in  a  direct  manner, 
and  that  the  effects  are  transmitted.  Thus,  as  is  generally  ad- 
mitted, the  European  settlers  in  the  United  States  undergo  a 
slight  but  extraordinarily  rapid  change  of  appearance.  Their 
bodies  and  limbs  become  elongated;  and  I  hear  from  Col.  Bernys 
that  during  the  late  war  in  the  United  States,  good  evidence  was 
afforded  of  this  fact  by  the  ridiculous  appearance  presented  by 
the  German  regiments,  when  dressed  in  ready-made  clothes 
manufactured  for  the  American  market,  and  which  were  much 
too  long  for  the  men  in  every  way.  There  is,  also,  a  considerable 
body  of  evidence  showing  that  in  the  Southern  States  the  house- 
slaves  of  the  third  generation  present  a  markedly  different  ap- 
pearance from  the  field-slaves.^® 

^  'Variation  of  Animals  and  Plants  under  Domestication,'  vol.  11. 
pp.  336,  337. 

^5  See,  for  instance,  Quatrefages  ('Revue  des  Cours  Scientifiques,' 
Oct.  10,  1868,  p.  724)  on  the  effects  of  residence  in  Abyssinia  and  Arabia, 
and  other  analogous  cases.  Dr.  Rolle  ('Der  Mensch,  seine  Abstam- 
mung,'  &c.,  1865,  s.  99)  states,  on  the  authority  of  Khanikof,  that  the 
greater  number  of  German  families  settled  in  Georgia,  have  acquired 
in  the  course  of  two  generations  dark  hair  and  eyes.  Mr.  D.  Forbes 
informs  me  that  the  Quichaus  in  the  Andes  vary  greatly  in  color, 
according  to  the  position  of  the  valleys  inhabited  by  them. 

88  Harlan,  'Medical  Researches,'  p.  532.  Quatrefages  ('Unite  de  I'Es- 
pece  Humaine.'  1861,  p.  128)  has  collected  much  evidence  on  this  head. 


192  THE  DESCENT  OF  MAN. 

If,  however,  we  look  to  the  races  of  man  as  distributed  over  the 
world,  we  must  infer  that  their  characteristic  differences  cannot 
be  accounted  for  by  the  direct  action  of  different  conditions  of 
life,  even  after  exposure  to  them  for  an  enormous  period  of  time. 
The  Esquimaux  live  exclusiv^ely  on  animal  food;  they  are 
clothed  in  thick  fur,  and  are  exposed  to  intense  cold  and  to  pro- 
longed darkness;  yet  they  do  not  differ  in  any  extreme  degree 
from  the  inhabitants  of  Southern  China,  who  live  entirely  on 
vegetable  food,  and  are  exposed  almost  naked  to  a  hot,  glaring 
climate.  The  unclothed  Fuegians  live  on  the  marine  productions 
of  their  inhospitable  shores;  the  Botocudos  of  Brazil  wander 
about  the  hot  forests  of  the  interior  and  live  chiefly  on  vegetable 
productions;  yet  these  tribes  resemble  each  other  so  closely  that 
the  Fuegians  on  board  the  "Beagle"  were  mistaken  by  some  Bra- 
zilians for  Botocudos.  The  Botocudos  again,  as  well  as  the  other 
inhabitants  of  tropical  America,  are  wholly  different  from  the 
Negroes  who  inhabit  the  opposite  shores  of  the  Atlantic,  a,re  ex- 
posed to  a  nearly  similar  climate,  and  follow  nearly  the  same 
habits  of  life. 

Nor  can  the  differences  between  the  races  of  man  be  accounted 
for  by  the  inherited  effects  of  the  increased  or  decreased  use  of 
parts,  except  to  a  quite  insignificant  degree.  Men  who  habitu- 
ally live  in  canoes,  may  have  their  legs  somewhat  stunted;  those 
who  inhabit  lofty  regions  may  have  their  chests  enlarged;  and 
those  who  constantly  use  certain  sense-organs  may  have  the 
cavities  in  which  they  are  lodged  somewhat  increased  in  size,  and 
their  features  consequently  a  little  modified.  With  civilized  na- 
tions, the  reduced  size  of  the  jaws  from  lessened  use — the  habit- 
ual play  of  different  muscles  serving  to  express  different  emo- 
tions—and the  increased  size  of  the  brain  from  greater  intellect- 
ual activity,  have  together  produced  a  considerable  effect  on 
their  general  appearance  when  compared  with  savages.®^  In- 
creased bodily  stature,  without  any  corresponding  increase  in 
the  size  of  the  brain,  may  (judging  from  the  previously  adduced 
case  of  rabbits),  have  given  to  some  races  an  elongated  skull  of 
the  dolichocephalic  type. 

Lastly,  the  little-understood  principle  of  correlated  develop- 
ment has  sometimes  come  into  action,  as  in  the  case  of  great 
muscular  development  and  strongly  projecting  supra-orbital 
ridges.  The  color  of  the  skin  and  hair  are  plainly  correlated,  as 
is  the  texture  of  the  hair  with  its  color  in  the  Mandans  of  North 
America.*^^    The  color  also  of  the  skin,  and  the  odor  emitted  by  it, 

07  See  Prof.  Schaaffhausen,  translat.  in  'Anthropological  Review,' 
Oct.   1868,   p.   429. 

63  Mr.  Catlin  states  ('N.  American  Indians,'  3rd  edit.  1842,  vol.  i.  p.  49) 
that  in  the  whole  trihe  of  the  Mandans,  about  one  in  ten  or  twelve 
of  the  members,  of  all  ages  and  both  sexes,  have  bright  silvery  gray 


THE  FORMATION  OF  RACES.  193 

are  likewise  in  some  manner  connected.  With  the  breeds  of  sheep 
the  number  of  hairs  within  a  given  space  and  the  number  of  the 
excretory  pores  are  related.^^  If  we  may  judge  from  the  analogy 
of  our  domesticated  animals,  many  modifications  of  structure  in 
man  probably  come  under  this  principle  of  correlated  develop- 
ment. 

We  have  now  seen  that  the  external  characteristic  differences 
between  the  races  of  man  cannot  be  accounted  for  in  a  satisfac- 
tory manner  by  the  direct  action  of  the  conditions  of  life,  nor  by 
the  effects  of  the  continued  use  of  parts,  nor  through  the  prin- 
ciple of  correlation.  We  are  therefore  led  to  inquire  whether 
slight  individual  differences,  to  which  man  is  eminently  liable, 
may  not  have  been  preserved  and  augmented  during  a  long  series 
of  generations  through  natural  selection.  But  here  we  are  at 
once  met  by  the  objection  that  beneficial  variations  alone  can  be 
thus  preserved;  and  as  far  as  we  are  enabled  to  judge,  although 
always  liable  to  err  on  this  head,  none  of  the  differences  between 
the  races  of  man  are  of  any  direct  or  special  service  to  him.  The 
intellectual  and  moral  or  social  faculties  must  of  course  be  ex- 
cepted from  this  remark.  The  great  variability  of  all  the  ex- 
ternal differences  between  the  races  of  man,  likewise  indicates 
that  they  cannot  be  of  much  importance;  for  if  important,  they 
would  long  ago  have  been  either  fixed  and  preserved,  or  elimi- 
nated. In  this  respect  man  resembles  those  forms,  called  by 
naturalists  protean  or  polymorphic,  which  have  remained  ex- 
tremely variable,  ov/ing,  as  it  seems,  to  such  variations  being  of 
an  indifferent  nature,  and  to  their  having  thus  escaped  the  ac- 
tion of  natural  selection. 

We  have  thus  far  been  baffled  in  all  our  attempts  to  account 
for  the  differences  between  the  races  of  man;  but  there  remains 
one  important  agency,  namely  Sexual  Selection,  which  appears  to 
have  acted  powerfully  on  man,  as  on  many  other  animals.  I  do 
not  intend  to  assert  that  sexual  selection  will  account  for  all  the 
differences  between  the  races.  An  unexplained  residuum  is  left, 
about  which  we  can  only  say,  in  our  ignorance,  that  as  individu- 
als are  continually  born  with,  for  instance,  heads  a  little  rounder 
or  narrower,  and  with  noses  a  little  longer  or  shorter,  such  slight 
differences  might  become  fixed  and  uniform,  if  the  unknown 
agencies  which  induced  them  were  to  act  in  a  more  constant 
manner,  aided  by  long-continued  intercrossing.  Such  variations 
come  under  the  provisional  class,  alluded  to  in  our  second  chap- 
hair,  which  is  hereditary.  Now  this  hair  is  as  coarse  and  harsh  as 
that  of  a  horse's  mane,  whilst  the  hair  of  other  colors  is  fine  and  soft. 
^»  On  the  odor  of  the  skin,  Godron,  'Sur  I'Espece,'  torn.  ii.  p.  217. 
On  the  pores  in  the  skin,  Dr.  Wilckens,  'Die  Aufgaben  der  Land- 
wirth.  Zootechnik,'  1869,  s.  1. 
14 


194  THE  DESCENT  OF  MAN. 

ter,  which  for  the  want  of  a  better  term  are  often  called  spon- 
taneous. Nor  do  I  pretend  that  the  efCects  of  sexual  selection  can 
be  indicated  with  scientific  precision;  but  it  can  be  shown  that 
it  would  be  an  inexplicable  fact  if  man  had  not  been  modified  by 
this  agency,  which  appears  to  have  acted  powerfully  on  innumer- 
able animals.  It  can  further  be  shown  that  the  differences  be- 
tween the  races  of  man,  as  in  color,  hairiness,  form  of  features, 
&c.,  are  of  a  kind  which  might  have  been  expected  to  come  under 
the  influence  of  sexual  selection.  But  in  order  to  treat  this  sub- 
ject properly,  I  have  found  it  necessary  to  pass  the  whole  animal 
kingdom  in  review.  I  have  therefore  devoted  to  it  the  Second 
Part  of  this  work.  At  the  close  I  shall  return  to  man,  and,  after 
attempting  to  show  how  far  he  has  been  modified  through  sexual 
selection,  will  give  a  brief  summary  of  the  chapters  in  this  First 
Part. 


Note  on  the  Resemblances  and  Diffeeences  in  the  Structuee 
AND  THE  Development  of  the  Beain  in  Man  and  Apes. 
By  Professoe  Huxley,   F.E.S. 

The  controversy  respecting  the  nature  and  the  extent  of  the 
differences  in  the  structure  of  the  brain  in  man  and  the  apes, 
which  arose  some  fifteen  years  ago,  has  not  yet  come  to  an  end, 
though  the  subject  matter  of  the  dispute  is,  at  present,  totallj'- 
different  from  what  it  was  formerly.  It  was  originally  asserted 
and  re-asserted,  with  singular  pertinacity,  that  the  brain  of  all 
apes,  even  the  highest,  differs  from  that  of  man,  in  the  absence  of 
such  conspicuous  structures  as  the  posterior  lobes  of  the  cerebral 
hemispheres,  with  the  posterior  cornu  of  the  lateral  ventricle  and 
the  hippocampus  minor,  contained  in  those  lobes,  which  are  so 
obvious  in  man. 

But  the  truth  that  the  three  structures  in  question  are  as  well 
developed  in  apes'  as  in  human  brains,  or  even  better;  and  that 
it  is  characteristic  of  all  the  Primates  (if  we  exclude  the  Lemurs) 
to  have  these  parts  well  developed,  stands  at  present  on  as  secure 
a  basis  as  any  proposition  in  comparative  anatomy.  Moreover,  it 
is  admitted  by  every  one  of  the  long  series  of  anatomists  who,  of 
late  years,  have  paid  special  attention  to  the  arrangement  of  the 
complicated  sulci  and  gyri  which  appear  upon  the  surface  of  the 
cerebral  hemispheres  in  man  and  the  higher  apes,  that  they  are 
disposed  after  the  very  same  pattern  in  him,  as  in  them.  Every 
principal  gyrus  and  sulcus  of  a  chimpanzee's  brain  is  clearly  rep- 
resented in  that  of  a  man,  so  that  the  terminology  which  applies 
to  the  one  answers  for  the  other.  On  this  point  there  is  no  dif- 
ference of  opinion.     Some  years  since.  Professor  Bischoff    pub- 


STRUCTURE    OF    THE    BRAIN.  195 

lished  a  memoir™  on  the  cerebral  convolutions  of  man  and  apes; 
and  as  the  purpose  of  my  learned  colleague  was  certainly  not  to 
diminish  the  value  of  the  differences  between  apes  and  men 
in  this  respect,  I  am  g'lad  to  make  a  citation  from  him. 

"That  the  apes,  and  especially  the  orang,  chimpanzee  and  gorilla, 
"come  very  close  to  man  in  their  organization,  much  nearer  than 
"to  any  other  animal,  is  a  well-known  fact,  disputed  by  nobody. 
"Looking  at  the  matter  from  the  point  of  view  of  organization 
"alone,  no  one  probably  would  ever  have  disputed  the  view  of 
"Linnaeus,  that  man  should  be  placed,  merely  as  a  peculiar  species, 
"at  the  head  of  the  mammalia  and  of  those  apes.  Both  show.  In 
"all  their  organs,  so  close  an  affinity,  that  the  most  exact  anatomi- 
"cal  investigation  is  needed  in  order  to  demonstrate  those  dif- 
"ferences  which  really  exist.  So  it  is  with  the  brains.  The  brains 
"of  man,  the  orang,  the  chimpanzee,  the  gorilla,  in  spite  of  all  the 
"important  differences  which  they  present,  come  very  close  to  one 
"another"  (1.  c.  p.  101). 

There  remains,  then,  no  dispute  as  to  the  resemblance  in  funda- 
mental characters,  between  the  ape's  brain  and  man's;  nor  any 
as  to  the  wonderfully  close  similarity  between  the  chimpanzee, 
orang  and  man,  in  even  the  details  of  the  arrangement  of  the 
gyri  and  sulci  of  the  cerebral  hemispheres.  Nor,  turning  to  the 
differences  between  the  brains  of  the  highest  apes  and  that  of  man, 
is  there  any  serious  question  as  to  the  nature  and  extent  of  these 
differences.  It  is  admitted  that  the  man's  cerebral  hemispheres 
are  absolutely  and  relatively  larger  than  those  of  the  orang  and 
chimpanzee;  that  his  frontal  lobes  are  less  excavated  by  the  up- 
ward protrusion  of  the  roof  of  the  orbils;  that  his  gyri  and  sulci 
are,  as  a  rule,  less  symmetrically  disposed,  and  present  a  greater 
number  of  secondary  plications.  And  it  is  admitted  that,  as  a  rule, 
in  man,  the  temporo-occipital  or  "external  perpendicular"  fissure, 
which  is  usually  so  strongly  marked  a  feature  of  the  ape's  brain 
is  but  faintly  marked.  But  it  is  also  clear,  that  none  of  these  dif- 
ferences constitutes  a  sharp  demarcation  between  the  man's  and 
the  ape's  brain.  In  respect  to  the  external  perpendicular  fissure 
of  Gratiolet,  in  the  human  brain,  for  instance.  Professor  Turner 
remarks  :^^ 

"In  some  brains  it  appears  simply  as  an  indentation  of  the  mar- 
"gin  of  the  hemispheres,  but,  in  others,  it  extends  for  some  distance 
"more  or  less  transversely  outwards.  I  saw  it  in  the  right  hemis- 
"phere  of  a  female  brain  pass  more  than  two  inches  outwards; 
"and  in  another  specimen,  also  the  right  hemisphere,  it  proceeded 

70  'Die  Grosshirn-Windungen  des  Menschen;'  'Abhandlung-en  der  K. 
Bayerischen  Akademie,'  Bd.  x.,  1868. 

■^  'Convolutions  of  the  Human  Cerebrum  Topographically  Consid- 
ered,' 1866,  j^  12. 


XS6  THE  DESCENT  OP  MAN. 

"for  four-tenths  of  an  inch  outwards,  and  then  extended  down- 
awards,  as  far  as  the  loAver  margin  of  the  outer  surface  of  the 
"hemisphere.  The  im.perfect  definition  of  this  fissure  in  the  ma- 
"jority  of  human  brains,  as  compared  with  its  remarkable  distinct- 
"ness  in  the  brain  of  most  Quadrumana,  is  owing  to  the  presence 
"in  the  former,  of  certain  superficial,  well  marked,  secondary  con- 
evolutions  which  bridge  it  over  and  connect  the  parietal  with  the 
"occipital  lobe.  The  closer  the  first  of  these  bridging  gyri  lies  to 
"the  longitudinal  fissure,  the  shorter  is  the  external  parieto-occip- 
"ital  fissure"  (1.  c.  p.  12). 

The  obliteration  of  the  external  perpendicular  fissure  of  Grat- 
iolet,  therefore,  is  not  a  constant  character  of  the  human  brain. 
On  the  other  hand,  its  full  development  is  not  a  constant  character 
of  the  higher  ape's  brain.  For,  in  the  chimpanzee,  the  more  or 
less  extensive  obliteration  of  the  external  perpendicular  sulcus  by 
"bridging  convolutions,"  on  one  side  or  the  other,  has  been  noted 
over  and  over  again  by  Prof.  Rolleston,  Mr.  Marshall,  M.  Broca 
and  Professor  Turner.  At  the  conclusion  of  a  special  paper  on 
this  subject  the  latter  writes :'^^ 

"The  three  specimens  of  the  brain  of  a  chimpanzee  just  de- 
"scribed,  prove,  that  the  generalization  which  Gratiolet  has  at- 
"tempted  to  draw  of  the  complete  absence  of  the  first  connecting 
"convolution  and  the  concealment  of  the  second,  as  essentially 
"characteristic  features  in  the  brain  of  this  animal,  is  by  no  means 
"universally  applicable.  In  only  one  specimen  did  the  brain,  in 
"these  particulars,  follow  the  law  which  Gratiolet  has  expressed. 
"As  regards  the  presence  of  the  superior  bridging-convolution,  I 
"am  inclined  to  think  that  it  has  existed  in  one  hemisphere,  at 
"least,  in  a  majority  6t  the  brains  of  this  animal  which  have,  up 
"to  this  time,  been  figured  or  described.  The  superficial  position 
"of  the  second  bridging  convolution  is  evidently  less  frequent,  and 
"has  as  yet,  I  believe,  only  been  seen  in  the  brain  (A)  recorded  in 
"this  communication.  The  asymmetrical  arrangement  in  the  con- 
"volutions  of  the  two  hemispheres,  v^^^hich  previous  observers  have 
"referred  to  in  their  descriptions  is  also  well  illustrated  in  these 
"specimens"  (pp.  8,  9). 

Even  were  the  presence  of  the  temporo-occipital,  or  external  per- 
pendicular, sulcus  a  mark  of  distinction  between  the  higher  apes 
and  man,  the  value  of  such  a  distinctive  character  would  be  ren- 
dered very  doubtful  by  the  structure  of  the  brain  in  the  Platy- 
rhine  apes.  In  fact  while  the  temporo-occipital  is  one  of  the  most 
constant  of  sulci  in  the  Catarhine,  or  Old  World,  apes,  it  is  never 
very  strongly  developed  in  the  New  World  apes;  it  is  absent  in 

"^^  Notes  more  especially  on  the  bridging-  convolutions  in  the  Brain 
of  the  Chimpanzee,  'Proceedings  of  the  Royal  Society  of  Edinburgh,' 
1865-6. 


STRUCTURE    OF    THE    BRAIN.  197 

the  smaller  Platyrhini;    rudimentary  in  Pithecia;"^  and  more  or 
less  obliterated  by  bridging  convolutions  in  Ateles. 

A  cliaracter  which  is  thus  variable  within  the  limits  of  a  single 
group  can  have  no  great  taxonomic  value. 

It  is  further  established,  that  the  degree  of  asymmetry  of  the 
convolution  of  the  two  sides  in  the  human  brain  is  subject  to 
much  individual  variation;  and  that,  in  those  individuals  of  the 
Bushman  race  who  have  been  examined,  the  gyri  and  sulci  of  the 
two  hemispheres  are  considerably  less  complicated  and  more  sym- 
metrical than  in  the  European  brain,  while,  in  some  individuals 
of  the  chimpanzee,  their  complexity  and  asymmetry  become  no- 
table. This  is  particularly  the  case  in  the  brain  of  a  young  male 
chimpanzee  figured  by  M.  Broca.  ('L'ordre  des  Primates,'  p.  165, 
fig.  11.) 

Again,  as  respects  the  question  of  absolute  size,  it  is.  estab- 
lished that  the  difference  between  the  largest  and  the  smallest 
healthy  human  brain  is  greater  than  the  difference  between  the 
smallest  healthy  human  brain  and  the  largest  chimpanzee's  or 
'  orang's  brain. 

Moreover,  there  is  one  circumstance  in  which  the  orang's  and 
chimpanzee's  brains  resemble  man's  but  in  which  they  differ  from 
the  lower  apes,  and  that  is  the  presence  of  two  corpora  candicantia 
— the  Cynomorpha  having  but  one. 

In  view  of  these  facts  I  do  not  hesitate  in  this  year  1874,  to  re- 
peat and  insist  upon  the  proposition  which  I  enunciated  in  186.S.'* 

"So  far  as  cerebral  structure  goes,  therefore,  it  is  clear  that  man 
"differs  less  from  the  chimpanzee  or  the  orang,  than  these  do  even 
"from  the  monkeys,  and  that  the  difference  between  the  brain  of 
"the  chimpanzee  and  of  man  is  almost  insignificant,  when  ccm- 
"pared  with  that  between  the  chimpanzee  brain  and  that  of  a  Le- 
*'mur." 

In  the  paper  to  which  I  have  referred.  Professor  Bischoff  does 
not  deny  the  second  part  of  this  statement,  but  he  first  makes  the 
irrelevant  remark  that  it  is  not  wonderful  if  the  brains  of  an  orang 
and  a  l^emur  are  very  different;  and  secondly,  goes  on  to  assert 
that,  "if  we  successively  compare  the  brain  of  a  man  with  that  of 
"an  orang;  the  brain  of  this  with  that  of  a  chimpanzee;  of  this 
"with  that  of  a  gorilla,  and  so  on  of  a  Hylobates,  Semnopithecus, 
•'Cynocephalus,  Cercopithecus,  Macacus,  Cebus,  Callithrix,  Lemur, 
"Stenops,  Hapale,  we  shall  not  meet  with  a  greater  or  even  as 
"great  a  break  in  the  degree  of  development  of  the  convolutions, 
"as  we  find  between  the  brain  of  a  man  and  that  of  an  orang  or 
"chimpanzee." 


^3  Flower   'On  the  Anatomy  of  Pithecia  Monachus,'    'Proceedings  of 
the  Zoological  Society,'  1862. 
■^^  'Man's  Place  in  Nature,'  p.  102. 
14 


198  THE   DESCENT  OF  MAN. 

To  wliich  I  reply,  firstly,  that  whether  this  assertion  be  true  o* 
false,  it  has  nothing  whatever  to  do  with  the  proposition  enun- 
ciated in  'Man's  Place  in  Nature,'  which  refers  not  to  the  develop- 
ment of  the  convolutions  alone,  but  to  the  structure  of  the  whole 
brain.  If  Professor  Bischoff  had  taken  the  trouble  to  refer  to  p. 
96  of  the  work  he  criticizes,  in  fact,  he  would  have  found  the  fol- 
lowing passage:  "And  it  is  a  remarkable  circumstance  that 
"though,  so  far  as  our  present  knowledge  extends,  there  is  one 
"true  structural  break  in  the  series  of  forms  of  Simian  brains, 
"this  hiatus  does  not  lie  between  man  and  the  manlike  apes,  but 
"between  the  lower  and  the  lowest  Simians,  or  in  other  words, 
"between  the  Old  and  New  World  apes  and  monkeys  and  the  Le- 
"murs.  Every  Lemur  which  has  yet  been  examined,  in  fact,  has 
''its  cerebellum  partially  visible  from  above;  and  its  posterior  lobe, 
"with  the  contained  posterior  cornu  and  hippocampus  minor,  more 
"or  less  rudimentary.  Every  marmoset,  American  monkey.  Old 
"World  monkey,  baboon,  or  manlike  ape,  on  the  contrary,  has  its 
"cerebellum  entirely  hidden,  posteriorly,  by  the  cerebral  lobes,  and 
"possesses  a  large  posterior  cornu  with  a  well-developed  hippo- 
"campus  minor." 

This  statement  was  a  strictly  accurate  account  of  what  was 
known  when  it  was  made;  and  it  does  not  appear  to  me  to  be  more 
than  apparently  weakened  by  the  subsequent  discovery  of  the 
relatively  small  development  of  the  posterior  lobes  in  the  Siamang 
and  in  the  Howling  monkey.  Notwithstanding  the  exceptional 
brevity  of  the  posterior  lobes  in  these  two  species,  no  one  will 
pretend  that  their  brains,  in  the  slightest  degree,  approach  those 
of  the  Lemurs.  And  if,  instead  of  putting  Hapale  out  of  its  nat- 
ural place,  as  Professor  BischofE  most  unaccountably  does,  we 
write  the  series  of  animals  he  has  chosen  to  mention  as  follows: 
Homo,  Pithecus,  Troglodytes,  Hylobates,  Semnopithecus,  Cynoce- 
phalus,  Cercopithecus,  Macacus,  Cebus,  Callithrix,  Hapale,  Lemur, 
Stenops,  I  venture  to  reaffirm  that  the  great  break  in  this  series 
lies  between  Hapale  and  Lemur,  and  that  this  break  is  consid- 
erably greater  than  that  between  any  other  two  terms  of  that 
series.  Professor  Bischoff  ignores  the  fact  that  long  before  he 
wrote,  Gratiolet  had  suggested  the  separation  of  the  Lemurs  from 
the  other  Primates  on  the  very  ground  of  the  difference  in  their 
cerebral  characters;  and  that  Professor  Flower  had  made  the  fol- 
lowing obervations  in  the  course  of  his  description  of  the  brain  of 
the  Javan  Loris.^^ 

"And  it  is  especially  remarkable  that,  in  the  development  of  the 
"posterior  lobes,  there  is  no  approximation  to  the  Lemurine,  short 
"hemisphered,  brain,  in  those  monkeys  which  are  commonly  sup- 


's 'Transactions  of  the  Zoological  Society,'  vol.  v.  1862. 


STRUCTURE  OF  THE  BRAIN.  199 

"posed  to  approach  this  family  in  other  respects,  viz.,  the  lower 
"members   of   the   Platyrhine   group." 

So  far  as  the  structure  of  the  adult  brain  is  concerned,  then,  the 
very  considerable  additions  to  our  knowledge,  which  have  been 
made  by  the  researches  of  so  many  investigators,  during  the  past 
ten  years,  fully  justify  the  statement  which  I  made  in  1863.  But 
it  has  been  said  that,  admitting  the  similarity  between  the  adult 
brains  of  man  and  apes,  they  are  nevertheless,  in  reality,  widely 
different,  because  they  exhibit  fundamental  differences  in  the 
mode  of  their  development.  No  one  would  be  more  ready  than 
I  to  admit  the  force  of  this  argument,  if  such  fundamental  dif- 
ferences of  development  really  exist.  But  I  deny  that  they  do 
exist.  On  the  contrary,  there  is  a  fundamental  agreement  in  the 
development  of  the  brain  in  men  and  apes. 

Gratiolet  originated  the  statement  that  there  is  a  fundamental 
difference  in  the  development  of  the  brains  of  apes  and  that  of 
man — consisting  of  this;  that,  in  the  apes,  the  sulci  which  first 
make  their  appearance  are  situated  on  the  posterior  region  of  the 
cerebral  hemispheres,  while,  in  the  human  foetus,  the  sulci  first  be- 
come visible  on  the  frontal  lobes.'® 

This  general  statement  is  based  upon  two  observations,  the  one 
of  a  Gibbon  almost  ready  to  be  born,  in  which  the  posterior  gyri 
were  "well  developed,"  while  those  of  the  frontal  lobes  were 
"hardly  indicated""^  (1.  c.  p.  39),  and  the  other  of  a  human  foetus 
at  the  22nd  or  23rd  week  of  uterogestation,  in  which  Gratiolet  notes 
that  the  insula  was  uncovered,  but  that  nevertheless  "des  incis- 
es "Chez  tous  les  singes,  les  plis  posterieurs  se  developpent  les  pre- 
"miers;  les  plis  anterieurs  se  developpent  plus  tard,  aussi  la  vertebre 
"occipitale  et  la  parietale  sont-elles  relativement  tresgrandes  chez  le 
"foetus  L'Homme  presente  une  exception  remarquable  quant  a 
"I'epoque  de  I'apparition  des  plis  frontaux,  qui  sont  les  premiers  in- 
"diques;  mais  le  developpement  general  du  lobe  frontal,  envisage 
"seulement  par  rapport  a  son  volume,  suit  les  memes  lois  que  dans 
"les  singes:"  Gratiolet,  'Memoire  sur  les  plis  cerebraux  de  THomme 
et  des  Primates,'  p.  39,  tab.  iv.  fig.  3. 

"  Gratiolet's  words  are  (1.  c.  p.  39):  "Dans  le  foetus  dont  il  s'agit 
"les  plis  cerebraux  posterieurs  sont  bien  developpes,  tandis  que  les 
"plis  du  lobe  frontal  sont  a  peine  indiques."  The  figure,  however 
(PI.  iv.  fig.  3),  shows  the  fissure  of  Rolando,  and  one  of  the  frontal 
sulci,  plainly  enough.  Nevertheless,  M.  Alix,  in  his  'Notice  sur  les 
travaux  anthropologiques  de  Gratiolet'  (Mem.  de  la  Societe  d'Anthro- 
pologie  de  Paris,  1868,  p.  32),  writes  thus:  "Gratiolet  a  eu  entre  les 
"mains  le  cerveau  d'un  foetus  de  Gibbon,  singe  eminemment  supe- 
"rieur,  et  tellement  rapproche  de  I'orang,  que  des  naturalistes  tres- 
"competents  I'ont  range  parmi  les  anthropoides.  M.  Huxley,  par  ex- 
"emple,  n'hesite  pas  sur  ce  point.  Eh  bien,  c'est  sur  le  cerveau  d'un 
"foetus  de  Gibbon  que  Gratiolet  a  vu  les  circonvolutions  du  lobe  tem- 
"poro-sphenoidal  deja  developpees  lorsqu'il  n'existent  pas  encore  de 


200  THE   DESCENT  OF  MAN. 

"ures  sement  le  lobe  antGrieur,  une  scissure  peu  profonde  indique 
"la  separation  dii  lobe  occipital,  tres-reduit  d'ailleurs  des  cette 
"epoque.  Le  reste  de  la  surface  cerebrale  est  encore  absolument 
"lisse." 

Three  views  of  the  brain  are  given  in  Plate  II.  figs.  1,  2,  3,  of 
the  work  cited,  showing  the  upper,  lateral  and  inferior  views  of 
the  hemispheres,  but  not  the  inner  view.  It  is  worthy  of  note 
that  the  figure  by  no  means  bears  out  Gratiolet's  description,  inas- 
much as  the  fissure  (anterotemporai)  on  the  posterior  half  of  the 
face  of  the  hemisphere  is  more  marked  than  any  of  those  vaguely 
indicated  in  the  anterior  half.  If  the  figure  is  correct  it  in  no  way 
justifies  Gratiolet's  conclusion:  "II  y  a  done  entre  ces  cerveaux 
"[those  of  a  Callithrix  and  of  a  Gibbon]  et  celui  du  foetus  humain 
"une  difference  fondamental.  Chez  celui-ci,  longtemps  avant  que 
"les  plis  temporaux  apparaissent,  les  plis  frontaux  essayent  d'ex- 
"ister." 

Since  Gratiolet's  time,  however,  the  development  of  the  gyri  and 
sulci  of  the  brain  has  been  made  the  subject  of  renewed  investiga- 
tion by  Schmidt,  Bischoff,  Pansch,^^  and  more  particularly  by 
Ecker,'^^  whose  work  is  not  only  the  latest,  but  by  far  the  most 
complete,  memoir  on  the  subject. 

The  final  results  of  their  inquiries  may  be  summed  up  as  fol- 
lows:— 

1.  In  the  human  foetus,  the  sylvian  fissure  is  formed  in  the 
course  of  the  third  month  of  uterogestation.  In  this,  and  in  the 
fourth  mouth,  the  cerebral  hemispheres  are  smooth  and  rounded 
(with  the  exception  of  the  sylvian  depression),  and  they  project 
backwards  far  beyond  the  cerebellum. 

2.  The  sulci,  properly  so  called,  begin  to  appear  in  the  interval 
between  the  end  of  the  fourth  and  the  beginning  of  the  sixth 
month  of  foetal  life,  but  Ecker  is  careful  to  point  out  that,  not 
only  the  time,  but  the  order,  of  their  appearance  is  subject  to  con- 
siderable individual  variation.  In  no  case,  however,  are  either 
the  frontal  or  the  temporal  sulci  the  earliest. 

The  first  which  appears,  in  fact,  lies  on  the  inner  face  of  the 
hemisphere  (whence  doubtless  Gratiolet,  who  does  not  seem  to 
have  examined  that  face  in  his  foetus,  overlooked  it),  and  is  either 

"plis  sur  le  lobe  frontal.  II  etait  done  bien  autorise  a  dire  que,  chez 
"I'honime  les  circonvolutions  apparaissent  d'CL  en  oo,  tandis  que  chez 
"les  singes  elles  se  developpent  d'cj  en  a." 

''^  'Ueber  die  typische  Anordnung  der  Furchen  und  Windung-en  auf 
den  Grosshirn-Hemispharen  des  Menschen  und  der  Affen.'  'Archiv 
fur  Anthropolog'ie,'  iii.,   1868. 

■^8  'Zur  Entwickelungs  Geschichte  der  Furchen  und  Windungen  der 
Grosshirn-Hemispharen  im  Foetus  des  Menschen.'  'Archiv  fur  An- 
thropologie,'  iii.,  1868. 


STRUCTURE    OF    THE    BRAIN.  201 

the  internal  perpendicular  (occipito-parietal),  or  the  calcarine  sul- 
cus, these  two  being  close  together  and  eventually  running  into 
one  another.  As  a  rule  the  occipito-parietal  is  the  earlier  of  the 
two. 

3.  At  the  latter  part  of  this  period,  another  sulcus,  the  "posterio, 
parietal,"  or  "Fissure  of  Rolando"  is  developed,  and  it  is  followed, 
in  the  course  of  the  sixth  month,  by  the  other  principal  sulci  of  the 
frontal,  parietal,  temporal  and  occipital  lobes.  There  is,  how- 
ever, no  clear  evidence  that  one  of  these  constantly  appears  be- 
fore the  other;  and  it  is  remarkable  that,  in  the  brain  at  the 
period  described  and  figured  by  Ecker  (1.  c.  p.  212-13,  Taf.  II.  figs,  1, 
2,  3,  4),  the  antero-temporal  sulcus  (scissure  parallele)  so  charac- 
teristic of  the  ape's  brain,  is  as  well,  if  not  better  developed  than 
the  fissure  of  Rolando,  and  is  much  more  marked  than  the  proper 
frontal  sulci. 

Taking  the  facts  as  they  now  stand,  it  appears  to  me  that  the 
order  of  the  appearance  of  the  sulci  and  gyri  in  the  fcetal  human 
brain  is  in  perfect  harmony  with  the  general  doctrine  of  evolu- 
tion, and  with  the  view  that  man  has  been  evolved  from  some 
ape-like  form;  though  there  can  be  no  doubt  that  that  form  was, 
in  many  respects,  different  from  any  member  of  the  Primates  now 
living. 

Von  Baer  taught  us,  half  a  century  ago,  that,  in  the  course  of 
their  development,  allied  animals  put  on,  at  first,  the  characters  of 
the  greater  groups  to  which  they  belong,  and,  by  degrees,  assume 
those  which  restrict  them  within  the  limits  of  their  family,  genus, 
and  species;  and  he  proved,  at  the  same  time,  that  no  develop- 
mental stage  of  a  higher  animal  is  precisely  similar  to  the  adult 
condition  of  any  lower  animal.  It  is  quite  correct  to  say  that  a 
frog  passes  through  the  condition  of  a  fish,  inasmuch  as  at  one 
period  of  its  life  the  tadpole  has  all  the  characters  of  a  fish,  and, 
if  it  went  no  further,  would  have  to  be  grouped  among  fishes. 
But  it  is  equally  true  that  a  tadpole  is  very  different  from  any 
known  fish. 

In  like  manner,  the  brain  of  a  human  foetus,  at  the  fifth  month, 
may  correctly  be  said  to  be,  not  only  the  brain  of  an  ape,  but  that 
of  an  Arctopithecine  or  marmoset-like  ape;  for  its  hemispheres, 
with  their  great  posterior  lobster,  and  with  no  sulci  but  the  syl- 
vian and  the  calcarine,  present  the  characteristics  found  only  in 
the  group  of  the  Arctopithecine  Primates,  But  it  is  equally  true, 
as  Gratiolet  remarks,  that,  in  its  widely  open  sylvian  fissure,  it 
differs  from  the  brain  of  any  actual  marmoset.  No  doubt  it  would 
be  much  more  similar  to  the  brain  of  an  advanced  foetus  of  a 
marmoset.  But  we  know  nothing  whatever  of  the  development  of 
the  brain  in  the  marmosets.  In  the  Platyrhini  proper,  the  only 
observation  with  which  I  am  acquainted  is  due  to  Pansch,  who 
found  in  the  brain  of  a  foetal  Cebus  Apella,  in  addition  to  the 


202  THE  DESCENT  OF  MAN. 

sylvian  fissure  and  the  deep  calcarine  fissure,  only  a  very  shallow 
anterotemporal  fissure  (scissure  parallele  of  Gratiolet). 

Now  this  fact,  taken  together  with  the  circumstance  that  the 
anterotemporal  sulcus  is  present  in  such  Platyrhini  as  the  Sai- 
miri,  which  present  mere  traces  of  sulci  on  the  anterior  half  of 
the  exterior  of  the  cerebral  hemispheres,  or  none  at  all,  undoubt- 
edly, so  far  as  it  goes,  affords  fair  evidence  in  favor  of  Gratiolet's 
hypothesis,  that  the  posterior  sulci  appear  before  the  anterior,  in 
the  brains  of  the  Platyrhini.  But,  it  by  no  means  follows,  that 
the  rule  which  may  hold  good  for  the  Platyrhini  extends  to  the 
Catarhini.  We  have  no  information  whatever  respecting  the  de- 
velopment of  the  brain  in  the  Cynomorpha;  and,  as  regards  the 
Anthropomorpha,  nothing  but  the  account  of  the  brain  of  the 
Gibbon,  near  birth,  already  referred  to.  At  the  present  moment, 
there  is  not  a  shadow  of  evidence  to  show  that  the  sulci  of  a  chim- 
panzee's, or  orang's,  brain  do  not  appear  in  the  same  order  as  a 
man's. 

Gratiolet  opens  his  preface  with  the  aphorism,  "II  est  dangereux 
"dans  les  sciences  de  conclure  trop  vite."  I  fear  he  must  have  for- 
gotten this  sound  maxim  by  the  time  he  had  reached  the  discus- 
sion of  the  differences  between  men  and  apes,  in  the  body  of  his 
work.  No  doubt,  the  excellent  author  of  one  of  the  most  re- 
markable contributions  to  the  just  understanding  of  the  mammal- 
ian brain  which  has  ever  been  made,  would  have  been  the  first 
to  admit  the  insufiiciency  of  his  data  had  he  lived  to  profit  by  the 
advance  of  inquiry.  The  misfortune  is  that  his  conclusions  have 
been  employed  by  persons  incompetent  to  appreciate  their  foun- 
dation, as  arguments  in  favor  of  obscurantism.^" 

But  it  is  important  to  remark  that,  whether  Gratiolet  was  right 
or  wrong  in  his  hypothesis  respecting  the  relative  order  of  appear- 
ance of  the  temporal  and  frontal  sulci,  the  fact  remains;  that,  be- 
fore either  temporal  or  frontal  sulci,  appear,  the  foetal  brain  of 
man  presents  characters  which  are  found  only  in  the  lowest  group 
of  the  Primates  (leaving  out  the  Lemurs) ;  and  that  this  is  ex- 
actly what  we  should  expect  to  be  the  case,  if  man  has  resulted 
from  the  gradual  modification  of  the  same  form  as  that  from 
which  the  other  Primates  have  sprung. 

^'^  For  example,  M.  I'Abbe  Lecomte  in  his  terrible  pamphlet  'Le  Dar- 
winisme  et  I'origine  de  I'Homme,'  1873. 


SEXUAL   SELECTION.  203 


PART  IL 

SEXUAL  SELECTION. 


CHAPTER  VIII. 
PRINCIPLES   OF   SEXUAL   SELECTION. 

Secondary  sexual  characters— Sexual  selection — Manner  of  action- 
Excess  of  males— Polygamy— The  male  alone  generally  modified 
through  sexual  selection— Eagerness  of  the  male — Variability  of 
the  male — Choice  exerted  by  the  female — Sexual  compared  with 
natural  selection — Inheritance,  at  corresponding  periods  of  life, 
at  corresponding  seasons  of  the  year,  and  as  limited  by  sex — Re- 
lations between  the  several  forms  of  inheritance — Causes  why  one 
sex  and  the  young  are  not  modified  through  sexual  selection— Sup- 
plement on  the  proportional  numbers  of  the  two  sexes  throughout 
the  animal  kingdom — The  proportion  of  the  sexes  in  relation  to 
natural  selection. 

With  animals  which  have  their  sexes  separated,  the  males 
necessarily  differ  from  the  females  in  their  organs  of  reproduc- 
tion; and  these  are  the  primary  sexual  characters.  But  the 
sexes  often  differ  in  what  Hunter  has  called  secondary  sexual 
characters,  which  are  not  directly  connected  with  the  act  of  re- 
production; for  instance,  the  male  possesses  certain  organs  of 
sense  or  locomotion,  of  which  the  female  is  quite  destitute,  or  has 
them  more  highly-developed,  in  order  that  he  may  readily  find  or 
reach  her;  or  again  the  male  has  special  organs  of  prehension 
for  holding  her  securely.  These  latter  organs,  of  infinitely  di- 
versified kinds,  graduate  into  those  which  are  commonly  ranked 
as  primary,  and  in  some  cases  can  hardly  be  distinguished  from 
them;  we  see  instances  of  this  in  the  complex  appendages  at  the 
apex  of  the  abdomen  in  male  insects.  Unless  indeed  we  confine 
the  term  "primary"  to  the  reproductive  glands,  it  is  scarcely 
possible  to  decide  v/hich  ought  to  be  called  primary  and  which 
secondary. 

The  female  often  differs  from  the  male  in  having  organs  for 


204  THE  DESCENT  OF  MAN. 

the  nourishment  or  protection  of  her  young,  such  as  the  mam- 
mary glands  of  mammals,  and  the  abdominal  sacks  of  the  marsu- 
pials. In  some  few  cases  also  the  male  possesses  similar  organs, 
which  are  wanting  in  the  female,  such  as  the  receptacles  for  the 
ova  in  certain  male  fishes,  and  those  temporarily  developed  in 
certain  male  frogs.  The  females  of  most  bees  are  provided  with 
a  special  apparatus  for  collecting  and  carrying  pollen,  and  their 
ovipositor  is  modified  into  a  sting  for  the  defense  of  the  larva3  and 
the  community.  Many  similar  cases  could  be  given,  but  they  do 
not  here  concern  us.  There  are,  however,  other  sexual  differ- 
ences quite  unconnected  with  the  primary  reproductive  organs, 
and  it  is  with  these  that  we  are  more  especially  concerned — such 
as  the  greater  size,  strength,  and  pugnacity  of  the  male,  his 
weapons  of  offense  or  means  of  defense  against  rivals,  his  gaudy 
coloring  and  various  ornaments,  his  power  of  song,  and  other 
such  characters. 

Besides  the  primary  and  secondary  sexual  differences,  such  as 
the  foregoing,  the  males  and  females  of  some  animals  differ  in 
structures  related  to  different  habits  of  life,  and  not  at  all,  or 
only  indirectly,  to  the  reproductive  functions.  Thus  the  females 
of  certain  flies  (Culicidse  and  Tabanidse)  are  blood-suckers, 
whilst  the  males,  living  on  flowers,  have  mouths  destitute  of 
mandibles.^  The  males  of  certain  moths  and  of  some  crustaceans 
(e.  g.  Tanais)  have  imperfect,  closed  mouths,  and  cannot  feed. 
The  complemental  males  of  certain  Cirripedes  live  like  epiphytic 
plants  either  on  the  female  or  the  hermaphrodite  form,  and  are 
destitute  of  a  mouth  and  of  prehensile  limbs.  In  these  cases  it  is 
the  male  which  has  been  modified,  and  has  lost  certain  important 
organs,  which  the  females  possess.  In  other  cases  it  is  the  fe- 
male which  has  lost  such  parts;  for  instance,  the  female  glow- 
worm is  destitute  of  wings,  as  also  are  many  female  moths,  some 
of  which  never  leave  their  cocoons.  Many  female  parasitic  crus- 
taceans have  lost  their  natatory  legs.  In  some  weevil-beetles 
(Curculionidae)  there  is  a  great  difference  betv\/^een  the  male  and 
female  in  the  length  of  the  rostrum  or  snout;-  but  the  meaning 
of  this  and  of  many  analogous  differences,  is  not  at  all  under- 
stood. Differences  of  structure  between  the  two  sexes  in  relation 
to  different  habits  of  life  are  generally  confined  to  the  lower  ani- 
mals; but  with  some  few  birds  the  beak  of  the  male  differs  from 
that  of  the  female.  In  the  Huia  of  New  Zealand  the  difference  is 
wonderfully  great,  and  we  hear  from  Dr.  Buller^  that  the  male 
uses  his  strong  beak  in  chiselling  the  larvae  of  insects  out  of  de- 

1  Westwood,  'Modern  Class,  of  Insects,'  vol.  ii.  1840,  p.  541.  For  the 
statement  about  Tanais,  mentioned  below,  I  am  indebted  to  Fritz 
Muller. 

2  Kirby  and  Spence,  'Introduction  to  Entomology,'  vol.  iii.  1826,  p.  309. 

3  'Birds  of  New  Zealand,'  1872,  p.  66. 


SEXUAL    SELECTION.  205 

cayed  wood,  whilst  the  female  probes  the  softer  parts  with  her 
far  longer,  much  curved  and  pliant  beak:  and  thus  they  mu- 
tually aid  ea^h  other.  In  most  cases,  differences  of  structure 
between  the  sexes  are  more  or  less  directly  connected  with  the 
propagation  of  the  species:  thus  a  female,  which  has  to  nourish 
a  multitude  of  ova,  requires  more  food  than  the  male,  and  con- 
sequently requires  special  means  for  procuring  it.  A  male  ani- 
mal, which  lives  for  a  very  short  time,  might  lose  its  organs  for 
procuring  food  through  disuse,  without  detriment;  but  he  would 
retain  his  locomotive  organs  in  a  perfect  state,  so  that  he  might 
reach  the  female.  The  female,  on  the  other  hand,  might  safely 
lose  her  organs  for  flying,  swimming,  or  walking,  if  she  gradu- 
ally acquired  habits  which  rendered  such  powers  useless. 

We  are,  however,  here  concerned  only  with  sexual  selection. 
This  depends  on  the  advantage  which  certain  individuals  have 
over  others  of  the  same  sex  and  species  solely  in  respect  of  repro- 
duction. When,  as  in  the  cases  above  mentioned,  the  two  sexes 
differ  in  structure  in  relation  to  different  habits  of  life,  they  have 
no  doubt  been  modified  through  natural  selection,  and  by  in- 
heritance limited  to  one  and  the  same  sex.  So  again  the  primary 
sexual  organs,  and  those  for  nourishing  or  protecting  the  young, 
come  under  the  same  influence;  for  those  individuals  which  gen- 
erated or  nourished  their  offspring  best,  would  leave,  casteris 
paribus,  the  greatest  number  to  inherit  their  superiority;  whilst 
those  which  generated  or  nourished  their  offspring  badly,  would 
leave  but  few  to  inherit  their  weaker  powers.  As  the  male  has 
to  flnd  the  female,  he  requires  organs  of  sense  and  locomotion, 
but  if  these  organs  are  necessary  for  the  other  purposes  of  life, 
as  is  generally  the  case,  they  will  have  been  developed  through 
natural  selection.  When  the  male  has  found  the  female,  he 
sometimes  absolutely  requires  prehensile  organs  to  hold  her; 
thus  Dr.  Wallace  informs  me  that  the  males  of  certain  moths 
cannot  unite  with  the  females  if  their  tarsi  or  feet  are  broken. 
The  males  of  many  oceanic  crustaceans,  when  adult,  have  their 
legs  and  antennas  modified  in  an  extraordinary  manner  for  the 
prehension  of  the  female;  hence  we  may  suspect  that  it  is  be- 
cause these  animals  are  washed  about  by  the  waves  of  the  open 
sea,  that  they  require  these  organs  in  order  to  propagate  their 
kind,  and  if  so,  their  development  has  been  the  result  of  ordinary 
or  natural  selection.  Some  animals  extremely  low  in  the  scale 
have  been  modified  for  this  same  purpose;  thus  the  males  of 
certain  parasitic  worms,  when  fully  grown,  have  the  lower  sur- 
face of  the  terminal  parts  of  their  bodies  roughened  like  a  rasp, 
and  with  this  they  coil  round  and  permanently  hold  the  females.* 

*  M.  Perrier  advances  this  case  ('Revue  Scientiflque/  Feb.  1,  1873, 
p.  865)  as  one  fatal  to  the  belief  in  sexual  selection,  inasmuch  as  he 
supposes   that  I   attribute   all    the   differences    between   the   sexes   to 


2561  THE  DESCENT  OF  MAN. 

When  the  two  sexes  follow  exactly  the  same  habits  of  life,  and 
the  male  has  the  sensory  or  locomotive  organs  more  highly  de- 
veloped than  those  of  the  female,  it  may  be  that  the  perfection  of 
these  is  indispensable  to  the  male  for  finding  the  female;  but  in 
the  vast  majority  of  cases,  they  serve  only  to  give  one  male  an 
advantage  over  another,  for  with  sufficient  time,  the  less  well- 
endowed  males  would  succeed  in  pairing  with  the  females;  and 
judging  from  the  structure  of  the  female,  they  would  be  in  all 
other  respects  equally  well  adapted  for  their  ordinary  habits  of 
life.  Since  in  such  cases  the  males  have  acquired  their  present 
structure,  not  from  being  better  fitted  to  survive  in  the  struggle 
for  existence,  but  from  having  gained  an  advantage  over  other 
males,  and  from  having  transmitted  this  advantage  to  their  male 
offspring  alone,  sexual  selection  must  here  have  come  into  action. 
It  was  the  importance  of  this  distinction  which  led  me  to  desig- 
nate this  form  of  selection  as  Sexual  Selection.  So  again,  if  the 
chief  service  rendered  to  the  male  by  his  prehensile  organs  is  to 
prevent  the  escape  of  the  female  before  the  arrival  of  other 
males,  or  when  assaulted  by  them,  these  organs  will  have  been 
perfected  through  sexual  selection,  that  is  by  the  advantage  ac- 
quired by  certain  individuals  over  their  rivals.  But  in  most  cases 
of  this  kind  it  is  impossible  to  distinguish  between  the  effects  of 
natural  and  sexual  selection.  Whole  chapters  could  be  filled 
with  details  on  the  differences  between  the  sexes  in  their  sensory, 
locomotive,  and  prehensile  organs.  As,  however,  these  struc- 
tures are  not  more  interesting  than  others  adapted  for  the  or- 
dinary purposes  of  life  I  shall  pass  them  over  almost  entirely, 
giving  only  a  few  instances  under  each  class. 

There  are  many  other  structures  and  instincts  which  must  have 
been  developed  through  sexual  selection — such  as  the  weapons 
of  offense  and  the  means  of  defense  of  the  males  for  fighting 
with  and  driving  away  their  rivals — their  courage  and  pugnacity 
— their  various  ornaments — their  contrivances  for  producing  vo- 
cal or  instrumental  music — and  their  glands  for  emitting  odors, 
most  of  these  latter  structures  serving  only  to  allure  or  excite 
the  female.  It  is  clear  that  these  characters  are  the  result  of 
sexual  and  not  of  ordinary  selection,  since  unarmed,  unorna- 
mented,  or  unattractive  males  would  succeed  equally  well  in  the 

sexual  selection.  This  disting-uished  naturalist,  therefore,  like  so 
many  other  Frenchmen,  has  not  taken  the  trouble  to  understand  even 
the  first  principles  of  sexual  selection.  An  English  naturalist  insists 
that  the  claspers  of  certain  male  animals  could  not  have  been  de- 
veloped throug-h  the  choice  of  tlie  female!  Had  I  not  met  with  this 
remark,  I  should  not  have  thought  it  possible  for  any  one  to  have 
read  this  chapter  and  to  have  imagined  that  I  maintain  that  the 
choice  of  the  female  had  anything  to  do  with  the  development  of  the 
prehensile  organs  in  the  male. 


SEXUAL    SELECTION.  207 

battle  for  life  and  in  leaving  a  numerous  progeny,  but  for  the 
presence  of  better  endowed  males.  We  may  infer  that  this 
would  be  the  case,  because  the  females,  which  are  unarmed  and 
unornamented,  are  able  to  survive  and  procreate  their  kind. 
Secondary  sexual  characters  of  the  kind  just  referred  to,  will  be 
fully  discussed  in  the  following  chapters,  as  being  in  many  re- 
spects interesting,  but  especially  as  depending  on  the  will,  choice, 
and  rivalry  of  the  individuals  of  either  sex.  When  we  behold 
two  males  fighting  for  the  possession  of  the  female,  or  several 
male  birds  displaying  their  gorgeous  plumage,  and  performing 
strange  antics  before  an  assembled  body  of  females,  we  cannot 
doubt  that,  though  led  by  instinct,  they  know  what  they  are 
about,  and  consciously  exert  their  mental  and  bodily  powers. 

Just  as  man  can  improve  the  breed  of  his  game-cocks  by  the 
selection  of  those  birds  which  are  victorious  in  the  cockpit,  so  it 
appears  that  the  strongest  and  most  vigorous  males,  or  those  pro- 
vided with  the  best  weapons,  have  prevailed  under  nature,  and 
have  led  to  the  improvement  of  the  natural  breed  or  species.  A 
slight  degree  of  variability  leading  to  some  advantage,  however 
slight,  in  reiterated  deadly  contests  would  suflBce  for  the  work  of 
sexual  selection;  and  it  is  certain  that  secondary  sexual  charac- 
ters are  eminently  variable.  Just  as  man  can  give  beauty,  ac- 
cording to  his  standard  of  taste,  to  his  male  poultry,  or  more 
strictly  can  modify  the  beauty  originally  acquired  by  the  parent 
species,  can  give  to  the  Sebright  bantam  a  new  and  elegant 
plumage,  an  erect  and  peculiar  carriage — so  it  appears  that  fe- 
male birds  in  a  state  of  nature,  have  by  a  long  selection  of  the 
more  attractive  males,  added  to  their  beauty  or  other  attractive 
qualities.  No  doubt  this  implies  powers  of  discrimination  and 
taste  on  the  part  of  the  female  which  will  at  first  appear  ex- 
tremely improbable;  but  by  the  facts  to  be  adduced  hereafter,  I 
hope  to  be  able  to  show  that  the  females  actually  have  these 
powers.  T^Hien,  however,  it  is  said  that  the  lower  animals  have 
a  sense  of  beauty,  it  must  not  be  supposed  that  such  sense  is 
comparable  with  that  of  a  cultivated  man,  with  his  multiform 
and  complex  associated  ideas.  A  more  just  comparison  would 
be  between  the  taste  for  the  beautiful  in  animals,  and  that  in  the 
lowest  savages,  who  admire  and  deck  themselves  with  any  bril- 
liant, glittering,  or  curious  object. 

From  our  ignorance  on  several  points,  the  precise  manner  in 
which  sexual  selection  acts  is  somewhat  uncertain.  Neverthe- 
less if  those  naturalists  who  already  believe  in  the  mutability  of 
species,  will  read  the  following  chapters,  they  will,  I  think,  agree 
with  me,  that  sexual  selection  has  played  an  important  part  in 
the  history  of  the  organic  world.  It  is  certain  that  amongst  al- 
most all  animals  there  is  a  struggle  between  the  males  for  the 
possession  of  the  female.    This  fact  is  so  notorious  that  it  would 


208  THE  DESCENT  OF  MAN. 

be  superfluous  to  give  instances.  Hence  the  females  have  the 
opportunity  of  selecting  one  out  of  several  males,  on  the  suppo- 
sition that  their  mental  capacity  suffices  for  the  exertion  of  a 
choice.  In  many  cases  special  circumstances  tend  to  make  the 
struggle  between  the  males  particularly  severe.  Thus  the  males 
of  our  migratory  birds  generally  arrive  at  their  places  of  breed- 
ing before  the  females,  so  that  many  males  are  ready  to  contend 
for  each  female.  I  am  informed  by  Mr.  Jenner  Weir,  that  the 
bird-catchers  assert  that  this  is  invariably  the  case  with  the 
nightingale  and  blackcap,  and  with  respect  to  the  latter  he  can 
himself  confirm  the  statement. 

Mr.  Swaysland  of  Brighton  has  been  in  the  habit,  during  the 
last  forty  years,  of  catching  our  migratory  birds  on  their  first 
arrival,  and  he  has  never  known  the  females  of  any  species  to 
arrive  before  their  males.  During  one  spring  he  shot  thirty-nine 
males  of  Ray's  wagtail  (Budytes  Raii)  before  he  saw  a  single  fe- 
male. Mr.  Gould  has  ascertained  by  the  dissection  of  those 
snipes  which  arrive  the  first  in  this  country,  that  the  males  come 
before  the  females.  And  the  like  holds  good  with  most  of  the 
migratory  birds  of  the  United  States.^  The  majority  of  the  male 
salmon  in  our  rivers,  on  coming  up  from  the  sea,  are  ready  to 
breed  before  the  females.  So  it  appears  to  be  with  frogs  and 
toads.  Throughout  the  great  class  of  insects  the  males  almost 
always  are  the  first  to  emerge  from  the  pupal  state,  so  that  they 
generally  abound  for  a  time  before  any  females  can  be  seen.® 
The  cause  of  this  difference  between  the  males  and  females  in 
their  periods  of  arrival  and  maturity  is  sufficiently  obvious. 
Those  males  which  annually  first  migrated  into  any  country,  or 
which  in  the  spring  were  first  ready  to  breed,  or  were  the  most 
eager,  would  leave  the  largest  number  of  offspring;  and  these 
would  tend  to  inherit  similar  instincts  and  constitutions.  It 
must  be  borne  in  mind  that  it  would  have  been  impossible  to 
change  very  materially  the  time  of  sexual  maturity  in  the  fe- 
males, without  at  the  same  time  interfering  with  the  period  of 
the  production  of  the  young — a  period  which  must  be  determined 
by  the  seasons  of  the  year.  On  the  whole  there  can  be  no  doubt 
that  with  almost  all  animals,  in  which  the  sexes  are  separate, 
there  is  a  constantly  recurrent  struggle  between  the  males  for  the 
possession  of  the  females. 

B  J,  A.  Allen,  on  the  'Mammals  and  Winter  Birds  of  Florida,'  Bull. 
Comp.   Zoology,  Harvard  College,   p.  268. 

«  Even  with  those  plants  in  which  the  sexes  are  separate,  the  male 
flowers  are  generally  mature  before  the  female.  As  first  shown  by 
C.  K.  Sprengel,  many  hermaphrodite  plants  are  dichogamous;  that 
is,  their  male  and  female  organs  are  not  ready  at  the  same  time,  so 
that  they  cannot  be  self-fertilized.  Now  in  such  flowers,  the  pol- 
len is  in  general  matured  before  the  stigma,  though  there  are  excep- 
tional cases  in  which  the  female  organs  are  beforehand. 


Sexual  selection.  209 

Our  difficulty  in  regard  to  sexual  selection  lies  in  understand- 
ing how  it  is  that  the  males  which  conquer  other  males,  or  those 
which  prove  the  most  attractive  to  the  females,  leave  a  greater 
number  of  offspring  to  inherit  their  superiority  than  their  beaten 
and  less  attractive  rivals.  Unless  this  result  does  follovf,  the 
characters  which  give  to  certain  males  an  advantage  over  others, 
could  not  be  perfected  and  augmented  through  sexual  selection. 
When  the  sexes  exist  in  exactly  equal  numbers,  the  worst-en- 
dowed males  will  (except  v/here  polygamy  prevails),  ultimately 
find  females,  and  leave  as  many  offspring,  as  well  fitted  for  their 
general  habits  of  life,  as  the  best-endowed  males.  From  various 
facts  and  considerations,  I  formerly  inferred  that  with  most  ani- 
mals, in  which  secondary  sexual  characters  are  well  developed, 
the  males  considerably  exceeded  the  females  in  number;  but  this 
is  not  by  any  means  always  true.  If  the  males  were  to  the  fe- 
males as  two  to  one,  or  as  three  to  two,  or  even  in  a  somewhat 
lower  ratio,  the  whole  affair  would  be  simple;  for  the  better- 
armed  or  more  attractive  males  would  leave  the  largest  number 
of  offspring.  But  after  investigating,  as  far  as  possible,  the  nu- 
merical proportion  of  the  sexes,  I  do  not  believe  that  any  great 
inequality  in  number  commonly  exists.  In  most  cases  sexua/1 
selection  appears  to  have  been  effective  in  the  following  manner. 

Let  us  take  any  species,  a  bird  for  instance,  and  divide  the  fe- 
males inhabiting  a  district  into  two  equal  bodies,  the  one  con- 
sisting of  the  more  vigorous  and  better-nourished  individuals 
and  the  other  of  the  less  vigorous  and  healthy.  The  former, 
there  can  be  little  doubt,  would  be  ready  to  breed  in  the  spring 
before  the  others;  and  this  is  the  opinion  of  Mr.  Jenner  Weir, 
who  has  carefully  attended  to  the  habits  of  birds  during  many 
years.  There  can  also  be  no  doubt  that  the  most  vigorous,  best- 
nourished  and  earliest  breeders  would  on  an  average  succeed  in 
rearing  the  largest  number  of  fine  offspring.''  The  males,  as  we 
have  seen,  are  generally  ready  to  breed  before  the  females;  the 
strongest,  and  with  some  species  the  best  armed  of  the  males, 
drive  away  the  weaker;  and  the  former  v/ould  then  unite  with 
the  more  vigorous  and  better-nourished  females,  because  they  are 
the  first  to   breed.^    Such   vigorous   pairs  would   surely  rear   a 

''  Here  is  excellent  evidence  on  the  character  of  the  offspring  from 
an  experienced  ornithologist.  Mr.  J.  A.  Allen,  in  speaking  ('Mam- 
mals and  Winter  Birds  of  E.  Florida,'  p.  229)  of  the  later  broods,  after 
the  accidental  destruction  of  the  first,  says,  that  these  "are  found  to 
"be  smaller  and  paler-colored  than  those  hatched  earlier  in  the  sea- 
"son.  In  cases  where  several  broods  are  reared  each  year,  as  a  gen- 
"eral  rule  the  birds  of  the  earlier  broods  seem  in  all  respects  the 
"most  perfect  and  vigorous." 

s  Hermann  Muller  has  come  to  this  same  conclusion  with  respect 
to  those  female  bees  which  are  the  first  to  emerge  from  the  pupa 
15 


210  THE  DESCENT  OF  MAN. 

larger  number  of  offspring  than  the  retarded  females,  which 
would  be  compelled  to  unite  with  the  conquered  and  less  power- 
ful males,  supposing  the  sexes  to  be  numerically  equal;  and  this 
is  all  that  is  wanted  to  add,  in  the  course  of  successive  genera- 
tions, to  the  size,  strength  and  courage  of  the  males,  or  to  im- 
prove their  weapons. 

But  in  very  many  cases  the  males  which  conquer  their  rivals, 
do  not  obtain  possession  of  the  females,  independently  of  the 
choice  of  the  latter.  The  courtship  of  animals  is  by  no  means  so 
simple  and  short  an  affair  as  might  be  thought.  The  females  are 
most  excited  by,  or  prefer  pairing  with,  the  more  ornamented 
males,  or  those  which  are  the  best  songsters,  or  play  the  best 
antics;  but  it  is  obviously  probable  that  they  would  at  the  same 
time  prefer  the  more  vigorous  and  lively  males,  and  this  has  in 
some  cases  been  cpnfirmed  by  actual  observation.^  Thus  the 
more  vigorous  females,  which  are  the  first  to  breed,  will  have  the 
choice  of  many  males;  and  though  they  may  not  always  select 
the  strongest  or  best  armed,  they  will  select  those  which  are 
vigorous  and  well  armed,  and  in  other  respects  the  most  at- 
tractive. Both  sexes,  therefore,  of  such  early  pairs  would,  as 
above  explained,  have  an  advantage  over  others  in  rearing  off- 
spring; and  this  apparently  has  sufficed  during  a  long  course  of 
generations  to  add  not  only  to  the  strength  and  fighting  powers 
of  the  males,  but  likewise  to  their  various  ornaments  or  other  at- 
tractions. 

In  the  converse  and  much  rarer  case  of  the  males  selecting  par- 
ticular females,  it  is  plain  that  those  which  were  the  most  vigor- 
ous and  had  conquered  others,  would  have  the  freest  choice;  and 
it  is  almost  certain  that  they  would  select  vigorous  as  well  as  at- 
tractive females.  Such  pairs  would  have  an  advantage  in  rear- 
ing offspring,  more  especially  if  the  male  had  the  power  to  de- 
fend the  female  during  the  pairing-season  as  occurs  with  some 
of  the  higher  animals,  or  aided  her  in  providing  for  the  young. 
The  same  principles  would  apply  if  each  sex  preferred  and  se- 
lected certain  individuals  of  the  opposite  sex;  supposing  that 
they  selected  not  only  the  more  attractive,  but  likewise  the  more 
vigorous  individuals. 

Nvcmerical  Proportion  of  the  Two  Sexes.  — I  have  remarked 
that  sexual  selection  would  be  a  simple  affair  if  the  males  were 
considerably  more  numerous  than  the  females.    Hence  I  was  led 


each  year.     See  his  remarkable  essay,  'Anwendung  der  Darwin'schen 
Lehre  auf  Bienen,'  'Verh.  d.  V.  Jahrg.'  xxix.  p.  45. 

»  With  respect  to  poultry,  I  have  received  information,  hereafter  to 
be  given,  to  this  effect.  Even  with  birds,  such  as  pigeons,  which 
pair  for  life,  the  female,  as  I  hear  from  Mr.  Jenner  Weir,  will  desert 
her  mate  if  he  is  injured  or  grows  weak. 


SEXUAL    SELECTION.  211 

to  investigate,  as  far  as  I  could,  the  proportions  between  the  two 
sexes  of  as  many  animals  as  possible;  but  the  materials  are 
scanty.  I  will  here  give  only  a  brief  abstract  of  the  results,  re- 
taining the  details  for  a  supplementary  discussion,  so  as  not  to 
interfere  with  the  course  of  my  argument.  Domesticated  ani- 
mals alone  afford  the  means  of  ascertaining  the  proportional 
numbers  at  birth;  but  no  records  have  been  specially  kept  for 
this  purpose.  By  indirect  means,  however,  I  have  collected  a  con- 
siderable body  of  statistics,  from  which  it  appears  that  with 
most  of  our  domestic  animals  the  sexes  are  nearly  equal  at  birth. 
Thus  25,560  births  of  race-horses  have  been  recorded  during 
twenty-one  years,  and  the  male  births  were  to  the  female  births 
as  99.7  to  100.  In  greyhounds  the  inequality  is  greater  than  with 
any  other  animal,  for  out  of  6878  births  during  twelve  years,  the 
male  births  were  to  the  female  as  110.1  to  100.  It  is,  however,  in 
some  degree  doubtful  whether  it  is  safe  to  infer  that  the  propor- 
tion would  be  the  same  under  natural  conditions  as  under  do- 
mestication; for  slight  and  unknown  differences  in  the  conditions 
affect  the  proportion  of  the  sexes.  Thus  with  mankind,  the  male 
births  in  England  are  as  104.5,  in  Russia  as  108.9,  and  with  the 
Jews  of  Livonia  as  120,  to  100  female  births.  But  I  shall  recur 
to  this  curious  point  of  the  excess  of  male  births  in  the  supple- 
ment to  this  chapter.  At  the  Cape  of  Good  Hope,  however,  male 
children  of  European  extraction  have  been  born  during  several 
years  in  the  proportion  of  between  90  and  99  to  100  female  chil- 
dren. 

For  our  present  purpose  we  are  concerned  with  the  proportion 
of  the  sexes,  not  only  at  birth,  but  also  at  maturity,  and  this  adds 
another  element  of  doubt;  for  it  is  a  well  ascertained  fact  that 
with  man  the  number  of  males  dying  before  or  during  birth,  and 
during  the  first  few  years  of  infancy,  is  considerably  larger  than 
that  of  females.  So  it  almost  certainly  is  with  male  lambs,  and 
probably  with  some  other  animals.  The  males  of  some  species 
kill  one  another  by  fighting;  or  they  drive  one  another  about 
until  they  become  greatly  emaciated.  They  must  also  be  often 
exposed  to  various  dangers,  whilst  wandering  about  in  eager 
search  for  the  females.  In  many  kinds  of  fish  the  males  are 
much  smaller  than  the  females,  and  they  are  believed  often  to  be 
devoured  by  the  latter,  or  by  other  fishes.  The  females  of  some 
birds  appear  to  die  earlier  than  the  males;  they  are  also  liable  to 
be  destroyed  on  their  nests,  or  whilst  in  charge  of  their  young. 
With  insects  the  female  larvae  are  often  larger  than  those  of  the 
males,  and  would  consequently  be  more  likely  to  be  de- 
voured. In  some  cases  the  mature  females  are  less  active  and 
less  rapid  in  their  movements  than  the  males,  and  could  not  es- 
cape so  well  from  danger.  Hence,  with  animals  in  a  state  of  na- 
ture, we  must  rely  on  mere  estimation,  in  order  to  judge  of  the 


212  THE  DESCENT  OF  MAN. 

proportions  of  the  sexes  at  maturity;  and  tliis  is  but  little  trust- 
worthy, except  when  the  inequality  is  strongly  marked.  Never- 
theless, as  far  as  a  judgment  can  be  formed,  we  may  conclude 
from  the  facts  given  in  the  supplement,  that  the  males  of  some 
few  mammals,  of  many  birds,  of  some  fish  and  insects,  are  con- 
siderably more  numerous  than  the  females. 

The  proportion  between  the  sexes  fluctuates  slightly  during 
successive  years:  thus  with  race-horses,  for  every  110  mares  born 
the  stallions  varied  from  107.1  in  one  year  to  92.6  in  another  year, 
and  with  greyhounds  from  116.3  to  95.3.  But  had  larger  num- 
bers been  tabulated  throughout  an  area  more  extensive  than  Eng- 
land, these  fluctuations  would  probably  have  disappeared;  and 
such  as  they  are,  would  hardly  suffice  to  lead  to  effective  sexual 
selection  in  a  state  of  nature.  Nevertheless,  in  the  cases  of 
some  few  wild  animals,  as  shown  in  the  supplement,  the  propor- 
tions seem  to  fluctuate  either  during  different  seasons  or  in  dif- 
ferent localities  in  a  sufficient  degree  to  lead  to  such  selection. 
For  it  should  be  observed  that  any  advantage,  gained  during  cer- 
tain years  or  in  certain  localities  by  those  males  which  were  able 
to  conquer  their  rivals,  or  were  the  most  attractive  to  the  females, 
would  probably  be  transmitted  to  the  offspring,  and  would  not 
subsequently  be  eliminated.  During  the  succeeding  seasons, 
when,  from  the  equality  of  the  sexes,  every  male  was  able  to  pro- 
cure a  female,  the  stronger  or  more  attractive  males  previously 
produced  would  still  have  at  least  as  good  a  chance  of  leaving  off- 
spring as  the  weaker  or  less  attractive. 

Polygamy. — The  practice  of  polygamy  leads  to  the  same  results 
as  would  follow  from  an  actual  inequality  in  the  number  of  the 
sexes;  for  if  each  male  secures  two  or  more  females,  many  males 
cannot  pair;  and  the  latter  assuredly  will  be  the  weaker  or  less 
attractive  individuals.  Many  mammals  and  some  few  birds  are 
polygamous,  but  with  animals  belonging  to  the  lower  classes  I 
have  found  no  evidence  of  this  habit.  The  intellectual  powers 
of  such  animals  are,  perhaps,  not  sufficient  to  lead  them  to  collect 
and  guard  a  harem  of  females.  That  some  relation  exists  be- 
tween polygamy  and  the  development  of  secondary  sexual  char- 
acters, appears  nearly  certain;  and  this  supports  the  view  that  a 
numerical  preponderance  of  males  would  be  eminently  favorable 
to  the  action  of  sexual  selection.  Nevertheless  many  animals, 
which  are  strictly  monogamous,  especially  birds,  display  strongly- 
marked  secondary  sexual  characters;  whilst  some  few  animals, 
which  are  polygamous,  do  not  have  such  characters. 

We  will  first  briefly  run  through  the  mammals,  and  then  turn 
to  birds.  The  gorilla  seems  to  be  polygamous,  and  the  male  dif- 
fers considerably  from  the  female;  so  it  is  with  some  baboons, 
which  live  in  herds  containing  twice  as  many  adult  females  as 


SEXUAL   SELECTION.  213 

males.  In  South  America  tlie  Mycetes  caraya  presents  well- 
marked  sexual  differences,  in  color,  beard,  and  vocal  organs;  and 
the  male  generally  lives  with  two  or  three  wives:  the  male  of  the 
Cebus  capucinus  differs  somewhat  from  the  female,  and  appears 
to  he  polygamous.^"  Little  is  known  on  this  head  with  respect 
to  most  other  monkeys,  but  some  species  are  strictly  monoga- 
mous. The  ruminants  are  eminently  polygamous,  and  they  pre- 
sent sexual  differences  more  frequently  than  almost  any  other 
group  of  mammals;  this  holds  good,  especially  in  their  weapons, 
but  also  in  other  characters.  Most  deer,  cattle,  and  sheep  are 
polygamous;  as  are  most  antelopes,  though  seme  are  monoga- 
mous. Sir  Andrew  Smith,  in  speaking  of  the  antelopes 
of  South  Africa,  says  that  in  herds  of  about  a  dozen  there  was 
rarely  more  than  one  m.ature  male.  The  Asiatic  Antilope  saiga 
appears  to  be  the  most  inordinate  polygamist  in  the  world;  for 
Pallas"  states  that  the  m.ale  drives  away  all  rivals,  and  collects  a 
herd  of  about  a  hundred  females  and  kids  together;  the  female 
is  hornless  and  has  softer  hair,  but  does  not  otherwise  differ  much 
from  the  male.  The  wild  horse  of  the  Falkland  Islands  and  of 
the  Western  States  of  North  America  is  polygamous,  but,  except 
in  his  greater  size  and  in  the  proportions  of  his  body,  differs  but 
little  from  the  mare.  The  wild  boar  presents  well-marked  sexual 
characters,  in  his  great  tusks  and  some  other  points.  In  Europe 
and  in  India  he  leads  a  solitary  life,  except  during  the  breeding- 
season;  but  as  is  believed  by  Sir  W.  Elliot,  who  has  had  many 
opportunities  in  India  of  observing  this  animal,  he  consorts  at 
this  season  with  several  females.  Whether  this  holds  good  in 
Europe  is  doubtful,  but  is  is  supported  by  some  evidence.  The 
adult  male  Indian  elephant,  like  the  boar,  passes  much  of  his 
time  in  solitude;  but  as  Dr.  Campbell  states,  when  with  others, 
"it  is  rare  to  find  more  than  one  male  with  a  whole  herd  of  fe- 
"males;"  the  larger  males  expelling  or  killing  the  smaller  and 
weaker  ones.  The  male  differs  from  the  female  in  his  immense 
tusks,  greater  size,  strength,  and  endurance;  so  great  is  the  dif- 
ference in  these  respects,  that  the  males  when  caught  are  valued 
at  one-fifth  more  than  the  females.^'    The  sexes  of  other  pachy- 


10  On  the  Gorilla,  Savage  and  Wyman.  'Boston  Journal  of  Nat.  Hist.' 
vol.  V.  1845-47,  p.  423.  On  Cynocephalus,  Brehm,  'Illust,  Thierleben,' 
B.  i.  1864,  s.  77.  On  Mycetes,  Rengger,  'Naturgesch. :  Saugethiere 
von  Paraguay,'  1830,  s.  14,  20.    Cebus,  Brehm,  ibid.  s.  108. 

"  Pallas,  'Spicilegia  Zoolog.,'  Pasc.  xii.  1777,  p.  29.  Sir  Andrew 
Smith,  'Illustrations  of  the  Zoology  of  S.  Africa,'  1849,  pi.  29,  on  the 
Kobus.  Owen,  in  his  'Anatomy  of  Vertebrates'  (vol.  iii.  1868,  p.  633) 
gives  a  table  shov/ing  incidentally  which  species  of  antelopes  are  gre- 
garious. 

^2  Dr.  Campbell,   in  'Proc.  Zoolog.  Soc'  1869,  p.   138.     See  also  an  in- 
teresting paper,   by  Lieut.   Johnstone,  in   'Proc.  Asiatic  Soc.   of  Ben- 
gal,' May,   1868. 
15 


214  THE  DESCENT  OF  MAN. 

dermatous  animals  differ  very  little  or  not  at  all,  and,  as  far  as 
known,  they  are  not  polygamists.  Nor  have  I  heard  of  any 
species  in  the  Orders  of  Cheiroptera,  Edentata,  Insectivora  and 
Rodents  being  polygamous,  excepting  that  amongst  the  Rodents, 
the  common  rat,  according  to  some  rat-catchers,  lives  with  several 
females.  Nevertheless  the  two  sexes  of  some  sloths  (Edentata) 
differ  in  the  character  and  color  of  certain  patches  of  hair  on 
their  shoulders."  And  many  kinds  of  bats  (Cheiroptera)  present 
well-marked  sexual  differences,  chiefly  in  the  males  possessing 
odoriferous  glands  and  pouches,  and  by  their  being  of  a  lighter 
color.^*  In  the  great  order  of  Rodents,  as  far  as  I  can  learn, 
the  sexes  rarely  differ,  and  when  they  do  so,  it  is  but  slightly  in 
the  tint  of  the  fur. 

As  I  heard  from  Sir  Andrew  Smith,  the  lion  in  South  Africa 
sometimes  lives  with  a  single  female,  but  generally  with  more, 
and,  in  one  case,  was  found  with  as  many  as  five  females;  so  that 
he  is  polygamous.  As  far  as  I  can  discover,  he  is  the  only  polyg- 
amist  amongst  all  the  terrestial  Carnivora,  and  he  alone  pre- 
sents well-marked  sexual  characters.  If,  however,  we  turn  to  the 
marine  Carnivora,  as  we  shall  hereafter  see,  the  case  is  widely 
different,  for  many  species  of  seals  offer  extraordinary  sexual 
differences,  and  they  are  eminently  polygamous.  Thus,  accord- 
ing to  Peron,  the  male  sea-elephant  of  the  Southern  Ocean  always 
possesses  several  females,  and  the  sea-lion  of  Porster  is  said  to  be 
surrounded  by  from  twenty  to  thirty  females.  In  the  North,  the 
male  sea-bear  of  Steller  is  accompanied  by  even  a  greater  number 
of  females.  It  is  an  interesting  fact,  as  Dr.  Gill  remarks,^'  that  in 
the  monogamous  species,  "or  those  living  in  small  communities, 
"there  is  little  difference  in  size  between  the  males  and  females; 
"in  the  social  species,  or  rather  those  of  which  the  males  have 
"harems,  the  males  are  vastly  larger  than  the  females." 

Amongst  birds,  many  species,  the  sexes  of  which  differ  greatly 
from  each  other,  are  certainly  monogamous.  In  Great  Britain 
we  see  well-marked  sexual  differences,  for  instance,  in  the  wild- 
duck  which  pairs  with  a  single  female,  the  common  blackbird, 
and  the  bullfinch  which  is  said  to  pair  for  life.  I  am  informed 
by  Mr.  Wallace  that  the  like  is  true  of  the  Chatterers  or  Cot- 
ingidse  of  South  America,  and  of  many  other  birds.  In  several 
groups  I  have  not  been  able  to  discover  whether  the  species  are 
polygamous  or  monogamous.  Lesson  says  that  birds  of  paradise, 
so  remarkable  for  their  sexual  differences,  are  polygamous,  but 
Mr.  Wallace  doubts  whether  he  had  sufficient  evidence.  Mr.  Sal- 
vin  tells  me  he  has  been  led  to  believe  that  humming-birds  are 
polygamous.    The  male  widow-bird,  remarkable  for  his  caudal 

"Dr.  Gray,  in  'Annals  and  Mag.  of  Nat.  Hist.'  1871,  p.  302. 

"  See  Dr.  Dobson's  excellent  paper,  in  'Proc.  Zoolog.  Soc'  1873,  p.  241. 

^5  The  Eared  Seals,    'American  Naturalist,'  vol.  iv.,  Jan.  1871. 


Sexual  selection.  215 

plumes,  certainly  seems  to  be  a  polygamist.^^  I  have  been  as- 
sured by  Mr.  Jenner  Weir  and  by  others,  that  it  is  somewhat 
common  for  three  starlings  to  frequent  the  same  nest;  but 
whether  this  is  a  case  of  polygamy  or  polyandry  has  not  been 
ascertained. 

The  Gallinacese  exhibit  almost  as  strongly  marked  sexual  dif- 
ferences as  birds  of  paradise  or  humming-birds,  and  many  of  the 
species  are,  as  is  well  known,  polygamous;  others  being  strictly 
monogamous.  ¥7hat  a  contrast  is  presented  between  the  sexes 
of  the  polygamous  peacock  or  pheasant,  and  the  monogamous 
guinea-fowl  or  partridge!  Many  similar  cases  could  be  given, 
as  in  the  grouse  tribe,  in  which  the  males  of  the  polygamous 
capercailzie  and  black-cock  differ  greatly  from  the  females;  whilst 
the  sexes  of  the  monogamous  red  grouse  and  ptarmigan  differ 
very  little.  In  the  Cursores,  except  amongst  the  bustards,  few 
species  offer  strongly-marked  sexual  differences,  and  the  great 
bustard  (Otis  tarda)  is  said  to  be  polygamous.  With  the  Gralla- 
tores,  extremely  few  species  differ  sexually,  but  the  ruff  (Machetes 
pugnax)  affords  a  marked  exception,  and  this  species  is  believed 
by  Montagu  to  be  a  polygamist.  Hence  it  appears  that  amongst 
birds  there  often  exists  a  close  relation  between  polygamy  and 
the  development  of  strongly-marked  sexual  differences.  I  asked 
Mr.  Bartlett,  of  the  Zoological  Gardens,  who  has  had  very  large 
experience  with  birds,  whether  the  male  tragopan  (one  of  the 
Gallinacese)  was  polygamous,  and  I  was  struck  by  his  answering, 
"I  do  not  knov/,  but  should  think  so  from  his  splendid  colors." 

It  deserves  notice  that  the  instinct  of  pairing  with  a  single  fe- 
male is  easily  lost  under  domestication.  The  wild-duck  is  strictly 
monogamus,  the  domestic-duck  highly  polygamous.  The  Rev. 
W.  D.  Fox  informs  me  that  out  of  some  half-tamed  wild-ducks, 
on  a  large  pond  in  his  neighborhood,  so  many  mallards  were  shot 
by  the  gamekeeper  that  only  one  was  left  for  every  seven  or 
eight  females;  yet  unusually  large  broods  were  reared.  The 
guinea-fowl  is  strictly  monogamous;  but  Mr.  Fox  finds  that  his 
birds  succeed  best  when  he  keeps  one  cock  to  tv/o  or  three  hens. 
Canary-birds  pair  in  a  state  of  nature,  but  the  breeders  in  Eng- 
land successfully  put  one  male  to  four  or  five  females.  I  have 
noticed  these  cases,  as  rendering  it  probable  that  wild  monog- 
amous species  might  readily  become  either  temporarily  or  per- 
manently polygamous. 

Too  little  is  known  of  the  habits  of  reptiles  and  fishes  to  en- 


18  'The  Ibis,'  vol.  iii.  1861,  p.  133,  on  the  Progne  Widow-bird.  See 
also  on  the  Vidua  axillaris,  ibid.  vol.  ii.  1860,  p.  211.  On  the  polyg-amy 
of  the  Capercailzie  and  Great  Bustard,  see  L.  Lloyd,  'Game  Birds  of 
Sweden,'  1867,  p.  19,  and  182.  Montag-u  and  Selby  speak  of  the  Black 
Grouse  as  polygamous  and  of  the  Red  Grouse  as  monogamous. 


216  THE  DESCENT  OF  MAN. 

able  us  to  speak  of  their  marriage  arrangements.  The  stickle- 
back (Gasterosteus),  however,  is  said  to  be  a  polygamist;'^  and 
the  male  during  the  breeding  season  differs  conspicuously  from 
the  female. 

To  sum  up  on  the  means  through  which,  as  far  as  we  can  judge, 
sexual  selection  has  led  to  the  development  of  secondary  sexual 
characters.  It  has  been  shown  that  the  largest  number  of  vig- 
orous offspring  will  be  reared  from  the  pairing  of  the  strongest 
and  best-armed  males,  victorious  in  contests  over  other  males, 
with  the  most  vigorous  and  best-nourished  females,  which  are 
the  first  to  breed  in  the  spring.  If  such  females  select  the  more 
attractive,  and  at  the  same  time  vigorous  males,  they  will  rear  a 
larger  number  of  offspring  than  the  retarded  females,  which  must 
pair  with  the  less  vigorous  and  less  attractive  males.  So  it  will 
be  if  the  more  vigorous  males  select  the  more  attractive  and  at 
the  same  time  healthy  and  vigorous  females;  and  this  will  es- 
pecially hold  good  if  the  male  defends  the  female,  and  aids  in  pro- 
viding food  for  the  young.  The  advantage  thus  gained  by  the 
more  vigorous  pairs  in  rearing  a  larger  number  of  offspring  has 
apparently  sufficed  to  render  sexual  selection  efficient.  But  a 
large  numerical  preponderance  of  males  over  females  will  be  still 
more  efficient;  whether  the  preponderance  is  only  occasional  and 
local,  or  permanent;  whether  it  occurs  at  birth,  or  afterwards 
from  the  greater  destruction  of  the  females;  or  whether  it  in- 
directly follows  from  the  practice  of  polygamy. 

The  Male  generally  more  modified  than  the  Female.  — Through- 
out the  animal  kingdom,  when  the  sexes  differ  in  external  appear- 
ance, it  is,  with  rare  exceptions,  the  male  which  has  been  the 
more  modified;  for,  generally,  the  female  retains  a  closer  resem- 
blance to  the  young  of  her  own  species,  and  to  other  adult  mem- 
bers of  the  same  group.  The  cause  of  this  seems  to  lie  in  the 
males  of  almost  all  animals  having  stronger  passions  than  the  fe- 
males. Hence  it  is  the  males  that  fight  together  and  sedulously 
display  their  charms  before  the  females;  and  the  victors  transmit 
their  superiority  to  their  male  offspring.  Why  both  sexes  do  not 
thus  acquire  the  characters  of  their  fathers,  will  be  considered 
hereafter.  That  the  males  of  all  mammals  eagerly  pursue  the  fe- 
males is  notorious  to  every  one.  So  it  is  with  birds;  but  many 
cock  birds  do  not  so  much  pursue  the  hen,  as  display  their  plum- 
age, perform  strange  antics,  and  pour  forth  their  song  in  her 
presence.  The  male  in  the  few  fish  observed  seems  much  more 
eager  than  the  female;  and  the  same  is  true  of  alligators,  and 
apparently  of  Batrachians.    Throughout  the  enormous  class  of  in- 

"  No€l   Humphreys,    'River  Gardens,'   1857. 


SEXUAL    SELECTION.  217 

sects,  as  Kirby  remarks/*  "the  law  is,  that  the  male  shall  seek 
"the  female."  Two  good  authorities,  Mr.  BlackwaTl  and  Mr.  C. 
Spence  Bate,  tell  me  that  the  males  of  spiders  and  crustaceans 
are  more  active  aiid  more  erratic  in  their  habits  than  the  female's. 
When  the  organs  of  sense  or  locomotion  are  present  in  the  one 
sex  of  insects  and  crustaceans  and  absent  in  the  other,  or  when, 
as  is  frequently  the  case,  they  are  more  highly  developed  in  the 
one  than  in  the  other,  it  is,  as  far  as  I  can  discover,  almost  in- 
variably the  male  which  retains  such  organs,  or  has  them  most 
developed;  and  this  shows  that  the  male  is  the  more  active  mem- 
ber in  the  courtship  of  the  sexes." 

The  female,  on  the  other  hand,  with  the  rarest  exceptions,  is 
less  eager  than  the  male.  As  the  illustrious  Hunter-°  long  ago 
observed,  she  generally  "requires  to  be  courted;"  she  is  coy,  and 
may  often  be  seen  endeavoring  for  a  long  time  to  escape  from  the 
male.  Every  observer  of  the  habits  of  animals  will  be  able  to  call 
to  mind  instances  of  this  kind.  It  is  shown  by  various  facts, 
given  hereafter,  and  by  the  results  fairly  attributable  to  sexual 
selection,  that  the  female,  though  comparatively  passive,  generally 
exerts  some  choice  and  accepts  one  male  in  preference  to  others. 
Or  she  may  accept,  as  appearances  would  sometimes  lead  us  to 
believe,  not  the  male  which  is  the  most  attractive  to  her,  but 
the  one  which  is  the  least  distasteful.  The  exertion  of  some 
choice  on  the  part  of  the  female  seems  a  law  almost  as  general 
as  the  eagerness  of  the  male. 

We  are  naturally  led  to  inquire  why  the  male,  in  so  many  and 
such  distinct  classes,  has  become  more  eager  than  the  female,  so 
that  he  searches  for  her,  and  plays  the  more  active  part  in  court- 
ship. It  would  be  no  advantage  and  some  loss  of  power  if  each 
sex  searched  for  the  other;  but  why  should  the  male  almost 
always  be  the  seeker?  The  ovules  of  plants  after  fertilization 
have  to  be  nourished  for  a  time;  hence  the  pollen  is  necessarily 
brought  to  the  female  organs — being  placed  on  the  stigma,  by 
means  of  insects  or  the  wind,  or  by  the  spontaneous  movements 

18  Kirby  and  Spence,  'Introduction  to  Entomology,'  vol.  iii.  1826,  p. 
342. 

19  One  parasitic  Hymenopterous  insect  (Westwood,  'Modern  Class, 
of  Insects,'  vol.  ii.  p.  160)  forms  an  exception  to  the  rule,  as  the  male 
has  rudimentary  Vv^ings,  and  never  quits  the  cell  in  which  it  is  born, 
whilst  the  female  has  well-developed  wing-s.  Audouin  believes  that 
the  females  of  this  species  are  impregnated  by  the  males  which  are 
born  in  the  same  cells  with  them;  but  it  is  much  more  probable  that 
the  females  visit  other  cells,  so  that  close  interbreeding  is  thus 
avoided.  We  shall  hereafter  meet  in  various  classes,  with  a  few  ex- 
ceptional cases,  in  which  the  female,  instead  of  the  male,  is  the 
seeker  and  wooer. 

20  'Essays  and  Observations,'  edited  by  Owen,  vol.  i.  1861,  p.  194, 


218  THE  DESCENT  OF  MAN. 

of  the  stamens;  and  in  the  Algse,  &c.,  by  the  locomotive  power 
of  the  antherozooids.  With  lowly-organized  aquatic  animals, 
permanently  affixed  to  the  same  spot  and  having  their  sexes  sep- 
arate, the  male  element  is  invariably  brought  to  the  female;  and 
of  this  we  can  see  the  reason,  for  even  if  the  ova  were  detached 
before  fertilization,  and  did  not  require  subsequent  nourishment  or 
protection,  there  would  yet  be  greater  difficulty  in  transporting 
them  than  the  male  element,  because  being  larger  than  the  latter, 
they  are  produced  in  far  smaller  numbers.  So  that  many  of  the 
lower  animals  are,  in  this  respect,  analogous  with  plants.^^  The 
male  of  affixed  and  aquatic  animals  having  been  led  to  emit 
their  fertilizing  element  in  this  way,  it  is  natural  that  any  of 
their  descendants,  which  rose  in  the  scale  and  became  locomotive, 
should  retain  the  same  habit;  and  they  would  approach  the  female 
as  closely  as  possible,  in  order  not  to  risk  the  loss  of  the  fertiliz- 
ing element  in  a  long  passage  of  it  through  the  water.  With  some 
few  of  the  lower  animals,  the  females  alone  are  fixed,  and  the 
males  of  these  must  be  the  seekers.  But  it  is  difficult  to  under- 
stand why  the  males  of  species,  of  which  the  progenitors  were 
primordially  free,  should  invariably  have  acquired  the  habit  of 
approaching  the  females,  instead  of  being  approached  by  them. 
But  in  all  cases,  in  order  that  the  males  should  seek  efficiently, 
it  would  be  necessary  that  they  should  be  endowed  with  strong 
passions;  and  the  acquirement  of  such  passions  would  naturally 
follow  from  the  more  eager  leaving  a  larger  number  of  offspring 
than  the  less  eager. 

The  great  eagerness  of  the  males  has  thus  indirectly  led  to  their 
much  more  frequently  developing  secondary  sexual  characters 
than  the  females.  But  the  development  of  such  characters  would 
be  much  aided,  if  the  males  were  more  liable  to  vary  than  the 
females— as  I  concluded  they  were— after  a  long  study  of  domes- 
ticated animals.  Von  Nathusius,  who  has  had  very  wide  experi- 
ence, is  strongly  of  the  same  opinion.^^  Good  evidence  also  in 
favor  of  this  conclusion  can  be  produced  by  a  comparison  of 
the  two  sexes  in  m.ankind.  During  the  Novara  Expedition^''  a 
vast  number  of  measurements  was  made  of  various  parts  of  the 
body  in  different  races,  and  the  men  were  found  in  almost  every 


21  Prof.  Sachs  C'Lehrbuch  der  Botanik,'  1870,  s.  633)  in  speaking  of 
the  male  and  female  reproductive  cells,  remarks,  "verhalt  sich  die 
"eine  bei  der  Vereinigung  activ,  ...  die  andere  erscheint  bei  der 
Vereinigung  passiv." 

22  'Vortrage    uber   Viehzucht,'   1872,    p.    63. 

23  'Reise  der  Novara:  Anthropolog-.  Theil,'  1867,  s.  216-269.  The  re- 
sults were  calculated  by  Dr.  Weisbach  from  measurements  made  by 
Drs.  K.  Scherzer  and  Schwarz.  On  the  greater  variability  of  the 
males  of  domesticated  animals,  see  my  'Variation  of  Animals  and 
Plants   under  Domestication,'  vol.   ii.  1868,   p.   75. 


SEXUAL.   SELECTION.  219 

case  to  present  a  greater  range  of  variation  than  the  women;    but 
I  shall  have  to  recur  to  this  subject  in  a  future  chapter.    Mr.  J. 
Wood,-^  who  has  carefullj^  attended  to  the  variation  of  the  muscles 
in  man,  puts  in  italics  the  conclusion  that  "the  greatest  number 
"of  abnormalities  in  each  subject  is  found  in  the  males."    He 
had  previously  remarked  that  "altogether  in  102  subjects,  the 
"varieties  of  redundancy  were  found  to  be  half  as  many  again  as 
"in  females,   contrasting  widely  with  the  greater  frequency  of 
"deficiency   in   females   before   described."    Professor  Macalister 
likewise  remarks-^  that  variations  in  the  muscles  "are  probably 
"more  common  in  males  than  females."    Certain  muscles,  which 
are  not  normally  present  in  mankind  are  also  more  frequently 
developed  in  the  male  than  in  the  female  sex,  although  exceptions 
to  this  rule  are  said  to  occur.    Dr.  Burt  Wilder-^  has  tabulated  the 
cases  of  152  individuals  with  supernumerary  digits,  of  which  86 
were  males,  and  89,  or  less  than  half,  females,  the  remaining  27 
being  of  unknown  sex.    It  should  not,  however,  be  overlooked 
that  women  would  more  frequently  endeavor  to  conceal  a  de- 
formity of  this  kind  than  men.    Again,  Dr.  L.  Meyer  asserts  that 
the  ears  of  man  are  more  variable  in  form  than  those  of  woman.^^ 
Lastly  the  temperature  is  more  variable  in  man  than  in  woman.^ 
The  cause  of  the  greater  general  variability  in  the  male  sex, 
than  in  the  female  is  unknown,  except  in  so  far  as  secondary 
sexual  characters  are  extraordinarily  variable,  and  are  usually 
confined  to  the  males;    and,  as  we  shall  presently  see,  this  fact  is, 
to  a  certain  extent,  intelligible.      Through  the  action  of  sexual 
and  natural  selection  male  animals  have  been  rendered  in  very 
many   instances   widely   different   from   their   females;     but   in- 
dependently of  selection  the  two  sexes,  from  differing  constitu- 
tionally, tend  to  vary  in  a  somewhat  different  manner.    The  fe- 
male has  to  expend  much  organic  matter  in  the  formation  of  her 
ova,  whereas  the  male  expends  much  force  in  fierce  contests  with 
his  rivals,  in  wandering  about  in  search  of  the  female,  in  exert- 
ing his  voice,  pouring  out  odoriferous  secretions,  &c.:    and  this 
expenditure  is  generally  concentrated  within  a  short  period.    The 
great  vigor  of  the  male  during  the  season  of  love  seems  often  to 
intensify    his    colors,    independently   of   any    marked    difference 
from  the  female.^    In  mankind,  and  even  as  lov/  down  in  the 


2^  'Proceeding's  Royal  See'   vol.   xvl.   July  1868,    pp.   519   and  524. 

25  'Proc.  Royal  Irish  Academy,'  vol.  x.  1868,  p.  123. 

-"  'Massachusetts  Medical   Soc'   vol.   ii.   No.   3,   1868,   p.   9. 

27  'Archiv  fur  Path.  Anat.   und  Phys.'  1871,  p.  488. 

^  The  conclusions  recently  arrived  at  by  Dr.  J.  Stockton  Hough, 
on  the  temperature  of  man,  are  given  in  the  'Pop.  Science  Review,' 
Jan.   1st,  1874  p.   97. 

29  Prof.  Manteg-azza  is  inclined  to  believe  ('Lettera  a  Carlo  Darwin,' 
'Archivip    per   I'Anthropologia,'    1871,    p.    306)    that   the   bright    colors, 


220  THE   DESCENT  OF  MAN. 

organic  scale  as  in  the  Lepidoptera,  the  temperature  of  the  body 
is  higher  in  the  male  than  in  the  female,  accompanied  in  the  case 
of  man  by  a  slower  pulse.^°  On  the  whole  the  expenditure  of 
matter  and  force  by  the  two  sexes  is  probably  nearly  equal,  though 
effected  in  very  different  ways  and  at  different  rates. 

From  the  causes  just  specified  the  tv/o  sexes  can  hardly  fail  to 
differ  somewhat  in  constitution,  at  least  during  the  breeding 
season:  and,  although  they  may  be  subjected  to  exactly  the  same 
conditions,  thej''  will  tend  to  vary  in  a  different  manner.  If  such 
variations  are  of  no  service  to  either  sex,  they  will  not  be  accu- 
mulated and  increased  by  sexual  or  natural  selection.  Neverthe- 
less, they  may  become  permanent  if  the  exciting  cause  acts  per- 
manently; and  in  accordance  with  a  frequent  form  of  inhsritance 
fhey  may  be  transmitted  to  that  sex  alone  in  vtrhich  they  first 
appeared.  In  this  case  the  two  sexes  will  come  to  present  perma- 
nent, yet  unimportant,  differences  of  character.  For  instance, 
Mr.  Allen  shows  that  with  a  large  number  of  birds  inhabiting  the 
northern  and  southern  United  States,  the  specimens  from  the 
south  are  darker-colored  than  those  from  the  north;  and  this 
seems  to  be  the  direct  result  of  the  difference  in  temperature, 
light,  ScQ,.,  between  the  two  regions.  Now,  in  some  few  cases,  the 
two  sexes  of  the  same  species  appear  to  have  been  differently  af- 
fected; in  the  Agelseus  phceniceus  the  males  have  had  their  colors 
greatly  intensified  in  the  south;  whereas  with  Cardinalis  virgin- 
ianus  it  is  the  females  which  have  been  thus  affected;  with  Quis- 
caius  major  the  females  have  been  rendered  extremely  variable  in 
tint,  whilst  the  males  remain  nearly  uniform.^^ 

A  fev/  exceptional  cases  occur  in  various  classes  of  animals,  in 
which  the  females  instead  of  the  males  have  acquired  well  pro- 
nounced secondary  sexual  characters,  such  as  brighter  colors, 
greater  size,  strength,  or  pugnacity.  With  birds  there  has  some- 
times been  a  complete  transposition  of  the  ordinary  characters 
proper  to  each  sex;  the  females  having  become  the  more  eager  in 
courtship,  the  males  remaining  comparatively  passive,  but  ap- 
parently selecting  the  more  attractive  females,  as  v/e  may  infer 
from  the  results.  Certain  hen  birds  have  thus  been  rendered 
more  highly  colored  or  otherwise  ornamented,  as  well  as  more 

common  in  so  many  male  animals,  are  due  to  the  presence  and  re- 
tention by  them  of  the  spermatic  fluid;  but  this  can  hardly  be  the 
case;  for  many  male  birds,  for  instance  young-  pheasants,  become 
brig^htly  colored   in   the   autumn  of  their   first  year. 

^^  For  mankind,  see  Dr.  J.  Stockton  Houg-h,  whose  conclusions  are 
given  in  the  'Pop.  Science  Review,'  1874,  p.  97.  See  Girard's  observa- 
tions on  the  Lepidoptera,  as  given  in  the  'Zoological  Record,'  1869,  p. 
347. 

31  'Mammals  and   Birds  of   E.    Florida,'  pp.  234,   280,   295. 


SEXUAL    SELECTION.  221 

powerful  and  pugnacious  than  the  cocks;    these  charact»ers  being 
transmitted  to  the  female  offspring  alone. 

It  may  be  suggested  that  in  some  cases  a  double  process  of  se^.; 
lection  has  been  carried  on;  that  the  males  have  selected  the' 
more  attractive  females,  and  the  latter  the  more  attractive  males. 
This  process,  however,  though  it  might  lead  to  the  modification  of  . 
both  sexes,  would  not  make  the  one  sex  different  from  the  other,-^^ 
unless  indeed  their  tastes  for  the  beautiful  differed;  but  this  is  a 
supposition  too  improbable  to  be  worth  considering  in  the  case  of 
any  animal,  excepting  man.  There  are,  however,  many  animals 
in  which  the  sexes  resemble  each  other,  both  being  furnished  with 
the  same  ornaments,  which  analogy  would  lead  us  to  attribute  to 
the  agency  of  sexual  selection.  In  such  cases  it  may  be  suggested 
with  more  plausibility,  that  there  has  been  a  double  or  mutual 
process  of  sexual  selection;  the  more  vigorous  and  precocious  fe- 
males selecting  the  more  attractive  and  vigorous  males,  the  latter 
rejecting  all  except  the  more  attractive  females.  But  from  what 
we  knov/  of  the  habits  of  animals,  this  view  is  hardly  probable, 
for  the  male  is  generally  eager  to  pair  with  any  female.  It  is 
more  probable  that  the  ornaments  common  to  both  sexes  were 
acquired  by  one  sex,  generally  the  male,  and  then  transmitted  to 
the  offspring  of  both  sexes.  If,  indeed,  during  a  lengthened  period 
the  males  of  any  species  were  greatly  to  exceed  the  females  in 
number,  and  then  during  another  lengthened  period,  but  under 
different  conditions,  the  reverse  were  to  occur,  a  double,  but  not 
simultaneous,  process  of  sexual  selection  might  easily  be  carried 
on,  by  which  the  two  sexes  might  be  rendered  widely  different. 

We  shall  hereafter  see  that  many  animals  exist,  of  which  neither 
sex  is  brilliantly  colored  or  provided  with  special  ornaments,  and 
yet  the  members  of  both  sexes  or  of  one  alone  have  probably  ac- 
quired simple  colors,  such  as  white  or  black,  through  sexual  selec- 
tion. The  absence  of  bright  tints  or  other  ornaments  may  be  the 
result  of  variations  of  the  right  kind  never  having  occurred,  or  of 
the  animals  themselves  having  preferred  plain  black  or  white. 
Obscure  tints  have  often  been  developed  through  natural  selection 
for  the  sake  of  protection,  and  the  acquirement  through  sexual 
selection  of  conspicuous  colors,  appears  to  have  been  sometimes 
checked  from  the  danger  thus  incurred.  But  in  other  cases  the 
males  during  long  ages  may  have  struggled  together  for  the  pos- 
session of  the  females,  and  yet  no  effect  v/ill  have  been  produced, 
unless  a  larger  number  of  offspring  were  left  by  the  more  success- 
ful males  to  inherit  their  superiority,  than  by  the  less  successful; 
and  this,  as  previously  shown,  depends  on  many  complex  contin- 
gencies. 

Sexual  selection  acts  in  a  less  rigorous  manner  than  natural  se- 
lection. The  latter  produces  its  effects  by  the  life  or  death  at  all 
ages  of  the  more  or  less  successful  individuals.    Death,  indeed. 


222  THE  DESCENT  OF  MAN. 

not  rarely  ensues  from  the  conflicts  of  rival  males.  But  generally 
the  less  successful  male  merely  fails  to  obtain  a  female,  or  obtains 
a  retarded  and  less  vigorous  female  later  in  the  season,  or,  if  polyg- 
amous, obtains  fewer  females;  so  that  they  leave  fewer,  less  vig- 
orous, or  no  offspring.  In  regard  to  structures  acquired  through 
ordinary  or  natural  selection,  there  is  in  most  cases,  as  long  as  the 
conditions  of  life  remain  the  same,  a  limit  to  the  amount  of  ad- 
vantageous modification  in  relation  to  certain  special  purposes; 
but  in  regard  to  structures  adapted  to  make  one  male  victorious 
over  another,  either  in  fighting  or  in  charming  the  female,  there 
is  no  definite  limit  to  the  amount  of  advantageous  modification; 
so  that  as  long  as  the  proper  variations  arise  the  work  of  sexual 
selection  will  go  on.  This  circumstance  may  partly  account  for 
tTie  frequent  and  extraordinary  amount  of  variability  presented 
by  secondary  sexual  characters.  Nevertheless,  natural  selection 
will  determine  that  such  characters  shall  not  be  acquired  by  the 
victorious  males,  if  they  would  be  highly  injurious,  either  by  ex- 
pending too  much  of  their  vital  powers,  or  by  exposing  them  to 
any  great  danger.  The  development,  however,  of  certain  struct- 
ures— of  the  horns,  for  instance,  in  certain  stags — has  been  car- 
ried to  a  wonderful  extreme;  and  in  some  cases  to  an  extreme 
which,  as  far  as  the  general  conditions  of  life  are  concerned,  must 
be  slightly  Injurious  to  the  male.  From  this  fact  we  learn  that  the 
advantages  which  favored  males  derive  from  conquering  other 
males  in  battle  or  courtship,  and  thus  leaving  a  numerous  prog- 
eny, are  in  the  long  run  greater  than  those  derived  from  rather 
more  perfect  adaptation  to  their  conditions  of  life.  We  shall  fur- 
ther see,  and  it  could  never  have  been  anticipated,  that  the  power 
to  charm  the  female  has  sometimes  been  more  important  than  the 
power  to  conquer  other  males  in  battle. 

LAWS   OF     INHERITANCE. 

In  order  to  understand  how  sexual  selection  has  acted  on  many 
animals  of  many  classes,  and  in  the  course  of  ages  has  produced 
a  conspicuous  result,  it  is  necessary  to  bear  in  mind  the  laws  of 
inheritance,  as  far  as  they  are  known.  Two  distinct  elements 
are  included  under  the  term  "inheritance"— the  transmission, 
and  the  development  of  characters;  but  as  these  generally  go 
together,  the  distinction  is  often  overlooked.  We  see  this  dis- 
tinction in  those  characters  which  are  transmitted  through  the 
early  years  of  life,  but  are  developed  only  at  maturity  or  during 
old  age.  We  see  the  same  distinction  more  clearly  with  secondary 
sexual  characters,  for  these  are  transmitted  through  both  sexes, 
though  developed  in  one  alone.  That  they  are  present  in  both 
sexes,  is  manifest  when  two  species,  having  strongly-marked  sex- 
ual characters,  are  crossed,  for  each  transmits  the  characters  prop- 


SEXUAL    SELECTION.  223 

er  to  its  own  male  and  female  sex  to  the  hybrid  offspring  of  either 
sex.  The  same  fact  is  likewise  manifest,  when  characters  proper 
to  the  male  are  occasionally  developed  in  the  female  when  she 
grows  old  or  becomes  diseased,  as,  for  instance,  when  the  com- 
mon hen  assumes  the  flowing  tail-feathers,  hackles,  comb,  spurs, 
voice,  and  even  pugnacity  of  the  cock.  Conversely,  the  same  thing 
is  evident,  more  or  less  plainly,  with  castrated  males.  Again, 
independently  of  old  age  or  disease,  characters  are  occasionally 
transferred  from  the  male  to  the  female,  as  when,  in  certain  breeds 
of  the  fowls,  spurs  regularly  appear  in  the  young  and  healthy  fe- 
males. But  in  truth  they  are  simply  developed  in  the  female;  for 
in  every  breed  each  detail  in  the  structure  of  the  spur  is  trans- 
mitted through  the  female  to  her  male  offspring.  Many  cases  will 
hereafter  be  given,  where  the  female  exhibits,  more  or  less  per- 
fectly, characters  proper  to  the  male,  in  whom  they  must  have 
been  first  developed,  and  then  transferred  to  the  female.  The 
converse  case  of  the  first  development  of  characters  in  the  female 
iind  of  transference  to  the  male,  is  less  frequent;  it  will  therefore 
be  well  to  give  one  striking  instance.  With  bees  the  pollen-col- 
lecting apparatus  is  used  by  the  female  alone  for  gathering  pollen 
for  the  larvae,  yet  in  most  of  the  species  it  is  partially  developed  in 
the  males  to  whom  it  is  quite  useless,  and  it  is  perfectly  developed 
in  the  males  of  Bombus  or  the  humble-bee.^^  As  not  a  single  other 
Hymenopterous  insect,  not  even  the  v/asp,  which  is  closely  allied 
to  the  bee,  is  provided  with  a  pollen-collecting  apparatus,  we  have 
no  grounds  for  supposing  that  male  bees  primordially  collected 
pollen  as  well  as  the  females;  although  we  have  some  reason  to 
suspect  that  male  mammals  primordially  suckled  their  young  as 
well  as  the  females.  Lastly,  in  all  cases  of  reversion,  characters 
are  transmitted  through  two,  three,  or  many  more  generations,  and 
are  then  developed  under  certain  unknown  favorable  conditions. 
This  important  distinction  between  transmission  and  development 
will  be  best  kept  in  mind  by  the  aid  of  the  hypothesis  of  pangen» 
esis.  According  to  this  hypothesis,  every  unit  or  cell  of  the  body 
throws  off  gemmules  or  undeveloped  atoms,  which  are  transmitted 
to  the  offspring  of  both  sexes,  and  are  multiplied  by  self-division. 
They  may  remain  undeveloped  during  the  early  years  of  life  or 
during  successive  generations;  and  their  development  into  units 
or  cells,  like  those  from  which  they  were  derived,  depends  on  their 
affinity  for,  and  union  with  other  units  or  cells  previously  devel- 
oped in  the  due  order  of  growth. 

Inheritance  at  corresponding  Periods  of  Life.  — This  tendency  is 
well  established.    A  new  character,  appearing  in  a  young  animal, 

'"•  M.  Muller,   'Anwendung  der  Darwin'schen  Lehre,'  &c.    Verh.  i.  n. 
■^     Jahrs,  xxix.  p.  42. 


224  '-THE  DESCENT  OF  MAN. 

whether  it  lasts  throughout  life  or  is  only  transient,  will,  in  gen- 
eral, reappear  in  the  offspring  at  the  same  age  and  last  for  the 
same  time.  If,  on  the  other  hand,  a  new  character  appears  at  ma- 
turity, or  even  during  old  age,  it  tends  to  reappear  in  the  offspring 
at  the  same  advanced  age.  When  deviations  from  this  rule  occur, 
the  transmitted  characters  much  oftener  appear  before,  than  after 
the  corresponding  age.  As  I  have  dwelt  on  this  subject  sufficiently 
in  another  workj^*^  I  will  here  merely  give  two  or  three  instances, 
for  the  sake  of  recalling  the  subject  to  the  reader's  mind.  In  sev- 
eral breeds  of  the  Fowl,  the  down-covered  chickens,  the  young 
bii-ds  in  their  first  true  plumage,  and  the  adults  differ  greatly  from 
one  another,  as  well  as  from  their  common  parent-form,  the  Gal- 
lus  bankiva;  and  these  characters  are  faithfully  transmitted  by 
each  breed  to  their  offspring  at  the  corresponding  periods  of  life. 
For  instance,  the  chickens  of  spangled  Hamburgs,  whilst  covered 
with  down,  have  a  few  dark  spots  on  the  head  and  rump,  but  are. 
not  striped  longitudinally,  as  in  many  other  breeds;  in  their  first 
true  plumage,  "they  are  beautifully  pencilled,"  that  is  each  feather 
is  transversely  marked  by  numerous  dark  bars;  but  in  their  second 
plumage  the  feathers  all  become  spangled  or  tipped  with  a  dark 
round  spot."*  Hence  in  this  breed  variations  have  occurred  at,  and 
been  transmitted  to,  three  distinct  periods  of  life.  The  Pigeon  of- 
fers a  more  remarkable  case,  because  the  aboriginal  parent  species 
does  not  undergo  any  change  of  plumage  with  advancing  age,  ex- 
cepting that  at  maturity  the  breast  becomes  more  iridescent;  yet 
there  are  breeds  which  do  not  acquire  their  characteristic  colors 
until  they  have  moulted  two,  three,  or  four  times;  and  these 
modifications  of  plumage  are  regularly  transmitted. 

InJieritance  at  corresponding  Seasons  of  the  Year. — With  ani- 
mals in  a  state  of  nature,  innumerable  instances  occur  of  charac- 
ters appearing  periodically  at  different  seasons.  We  see  this  in 
the  horns  of  the  stag,  and  in  the  fur  of  arctic  animals  which  be- 
comes  thick  and  white  during  the  winter.  Many  birds  acquire 
bright  colors  and  other  decorations  during  the  breeding-season 
alone.    Pallas  states,^^  that  in  Siberia  domestic  cattle  and  horses 

33  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
ii.  1868,  p.  75.  In  the  last  chapter  but  one,  the  provisional  hypothesis 
of  pangenesis,  above  alluded  to,  is  fully  explained. 

3*  These  facts  are  given  on  the  high  authority  of  a  great  breeder, 
Mr.  Teebay;  see  Tegetmeier's  'Poultry  Book,'  1868,  p.  158.  On  the 
characters  of  chickens  of  different  breeds,  and  on  the  breeds  of  the 
pigeon,  alluded  to  in  the  following  paragraph,  see  'Variation  of  Ani- 
mals,' &c.,  vol.  i.  pp.  160,  249;    vol.  ii.  p.  77. 

35  'Novae  species  Quadrupedum  e  Glirium  ordine,'  1778,  p.  7.  On 
the  transmission  of  color  by  the  horse,  see  'Variation  of  Animals, 
&c.,  under  Domestication,'  vol.  i.  p.  51.  Also  vol.  ii.  p.  71,  for  a  gen- 
eral discussion  on   'Inheritance  as  limited  by  Sex.' 


SEXUAL    SELECTION.  225 

become  lighter-colored  during  the  winter;  and  I  have  myself  ob- 
served, and  heard  of  similar  strongly  marked  changes  of  color, 
that  is,  from  brownish  cream-color  or  reddish-brown  to  a  perfect 
white  in  several  ponies  in  England.  Although  I  do  not  know  that 
this  tendency  to  change  the  color  of  the  coat  during  different  sea- 
sons is  transmitted,  yet  it  probably  is  so,  as  all  shades  of  color  are 
strongly  inherited  by  the  horse.  Nor  is  this  form  of  inheritance, 
as  limited  by  the  seasons,  more  remarkable  than  its  limitation  by 
age  or  sex. 

Inheritance  as  Limited  by  Sex. — The  equal  transmission  of 
characters  to  both  sexes  is  the  commonest  form  of  inheritance,  at 
least  with  those  animals  which  do  not  present  strongly-marked 
sexual  differences,  and  indeed  with  many  of  these.  But  characters 
are  somewhat  commonly  transferred  exclusively  to  that  sex,  in 
which  they  first  appear.  Ample  evidence  on  this  head  has  been 
advanced  in  my  work  on  'Variation  under  Domestication,'  but  a 
few  instances  may  here  be  given.  There  are  breeds  of  the  sheep 
and  goat,  in  which  the  horns  of  the  male  differ  greatly  in  shape 
from  those  of  the  female;  and  these  differences,  acquired  under 
domestication,  are  regularly  transmitted  to  the  same  sex.  As  a 
rule,  it  is  the  females  alone  in  cats  which  are  tortoise-shell,  the 
corresponding  color  in  the  males  being  rusty-red.  With  most 
breeds  of  the  fowl,  the  characters  proper  to  each  sex  are  trans- 
mitted to  the  same  sex  alone.  So  general  is  this  form  of  transmis- 
sion that  it  is  an  anomaly  when  variations  in  certain  breeds  are 
transmitted  equally  to  both  sexes.  There  are  also  certain  sub- 
breeds  of  the  fowl  in  which  the  males  can  hardly  be  distinguished 
from  one  another,  whilst  the  females  differ  considerably  in  color. 
The  sexes  of  the  pigeon  in  the  parent-species  do  not  differ  in  any 
external  character;  nevertheless,  in  certain  domesticated  breeds 
the  male  is  colored  differently  from  the  female.^"  The  wattle  in  the 
English  Carrier  pigeon,  and  the  crop  in  the  Pouter,  are  more 
highly  developed  in  the  male  than  in  the  female;  and  although 
these  characters  have  been  gained  through  long-continued  selec- 
tion by  man,  the  slight  differences  between  the  sexes  are  wholly 
due  to  the  form  of  inheritance  which  has  prevailed;  for  they  have 
arisen,  not  from,  but  rather  in  opposition  to,  the  wish  of  the 
breeder. 

Most  of  our  domestic  races  have  been  formed  by  the  accumula- 
tion of  many  slight  variations:  and  as  some  of  the  successive  steps 
have  been  transmitted  to  one  sex  alone,  and  some  to  both  sexes,  we 
find  in  the  different  breeds  of  the  same  species  all  gradations  be- 

88  Dr.  Chapuis,    'Le  Pigeon  "Voyageur  Beige,'  1865,  p.  87.     Boitard  et 
Corbie,   'Les  Pigeons  de  Voliere,'   &c.,  1824,  p.   173.    See,   also,   on  sim- 
ilar differences  in  certain  breeds  at  Modena,    'Le  variazioni  del  Col- 
ombi   domestici,'   del  Paolo  Bonizzi,  1873. 
16 


226  THE  DESCENT  OP  MAN. 

tween  great  sexual  dissimilarity  and  complete  similarity.  In- 
stances have  already  been  given  with  the  breeds  of  the  fowl  and 
pigeon,  and  under  nature  analogous  cases  are  common.  With 
animals  under  domestication,  but  whether  in  nature  I  will  not 
venture  to  say,  one  sex  may  lose  characters  proper  to  it,  and  may 
thus  come  somewhat  to  resemble  the  opposite  sex;  for  instance, 
the  males  of  some  breeds  of  the  fowl  have  lost  their  masculine  tail- 
plumes  and  hackles.  On  the  other  hand,  the  differences  betv/een 
the  sexes  may  be  increased  under  domestication,  as  with  merino 
sheep,  in  which  the  ewes  have  lost  their  horns.  Again,  characters 
proper  to  one  sex  may  suddenly  appear  in  the  other  sex;  as  in  those 
sub-breeds  of  the  fowl  in  which  the  hens  acquire  spurs  whilst 
young;  or,  as  in  certain  Polish  sub-breeds,  in  which  the  females, 
as  there  is  reason  to  believe,  originally  acquired  a  crest,  and  sub- 
sequently transferred  it  to  the  males.  All  these  cases  are  intelli- 
gible on  the  hypothesis  of  pangenesis;  for  they  depend  on  the 
gemmules  of  certain  parts,  although  present  in  both  sexes,  becom- 
ing, through  the  influence  of  domestication,  either  dormant  or  de- 
veloped in  either  sex. 

There  is  one  difficult  question  which  it  will  be  convenient  to 
defer  to  a  future  chapter;  namely,  whether  a  character  at  first  de- 
veloped in  both  sexes,  could  through  selection  be  limited  in  its  de- 
velopment to  one  sex  alone.  If,  for  instance,  a  breeder  observed 
that  some  of  his  pigeons  (of  which  the  characters  are  usually 
transferred  in  an  equal  degree  to  both  sexes)  varied  into  pale  blue, 
could  he  by  long-continued  selection  make  a  breed  in  which  the 
males  alone  should  be  of  this  tint,  whilst  the  females  remained 
unchanged?  I  will  here  only  say,  that  this,  though  perhaps  not 
impossible,  would  be  extremely  difficult;  for  the  natural  result  of 
breeding  from  the  pale-blue  males  would  be  to  change  the  whole 
stock  of  both  sexes  to  this  tint.  If,  however,  variations  of  the  de- 
sired tint  appeared,  which  were  from  the  first  limited  in  their 
development  to  the  male  sex,  there  would  not  be  the  least  dif- 
ficulty in  making  a  breed  with  the  two  sexes  of  a  different  color, 
as  indeed  has  been  effected  with  a  Belgian  breed,  in  which  the 
males  alone  are  streaked  with  black.  In  a  similar  manner,  if  any 
variation  appeared  in  a  female  pigeon,  which  was  from  the  first 
sexually  limited  in  its  development  to  the  females,  it  would  be 
easy  to  make  a  breed  with  the  females  alone  thus  characterized; 
but  if  the  variation  was  not  thus  originally  limited,  the  process 
would  be  extremely  difficult,  perhaps  impossible." 

3'^  Since  the  publication  of  the  first  edition  of  this  work,  it  has  been 
highly  satisfactory  to  me  to  find  the  following  remarks  (the  'Field,' 
Sept.  1872)  from  so  experienced  a  breeder  as  Mr.  Tegetmeier.  After 
describing  some  curious  cases  in  pigeons,  of  the  transmission  of  color 
by  one  sex  alone,  and  the  formation  of  a  sub-breed  with  this  char- 
acter,   he   says:     "It    is    a   singular    circumstance   that   Mr.    Darwin 


SEXUAL    SELECTION.  227 


On  the  Relatimi  between  the  Period  of  Development  of  a  Character 
and  its  Transmission  to  one  Sex  or  to  both  Sexes,  — Why  cer- 
tain characters  should  be  inherited  by  both  sexes,  and  other  char- 
acters by  one  sex  alone,  namely  by  that  sex  in  which  the  character 
first  appeared,  is  in  most  cases  quite  unknown.  We  cannot  even 
conjecture  why  with  certain  sub-breeds  of  the  pigeon,  black  striae, 
though  transmitted  through  the  female,  should  be  developed  in  the 
male  alone,  whilst  every  other  character  is  equally  transferred  to 
both  sexes.  Why,  again,  with  cats,  the  tortoise-shell  color  should, 
with  rare  exceptions,  be  developed  in  the  female  alone,  't'he  very 
same  character,  such  as  deficient  or  supernumerary  digits,  color- 
blindness, &c.,  may  with  mankind  be  inherited  by  the  males  alone 
of  one  family,  and  in  another  family  by  the  females  alone,  though 
in  both  cases  transmitted  through  the  opposite  as  well  as  through 
the  same  sex.^®  Although  we  are  thus  ignorant,  the  two  following 
rules  seem  often  to  hold  good — that  variations  which  first  appear 
in  either  sex  at  a  late  period  of  life,  tend  to  be  developed  in  the 
same  sex  alone;  whilst  variations  vv^hich  first  appear  early  in  life 
in  either  sex  tend  to  be  developed  in  both  sexes.  I  am,  however, 
far  from  supposing  that  this  is  the  sole  determining  cause.  As  I 
have  not  elsewhere  discussed  this  subject,  and  as  it  has  an  impor- 
tant bearing  on  sexual  selection,  I  must  here  enter  into  lengthy 
and  somewhat  intricate  details. 

It  is  in  itself  probable  that  any  character  appearing  at  an  early 
age  would  tend  to  be  inherited  equally  by  both  sexes,  for  the  sexes 
do  not  differ  much  in  constitution  before  the  power  of  reproduc- 
tion is  gained.  On  the  other  hand,  after  this  power  has  been 
gained  and  the  sexes  have  come  to  differ  in  constitution,  the  gem- 
mules  (if  I  may  again  use  the  language  of  pangenesis)  which  are 
cast  off  from  each  varying  part  in  the  one  sex  would  be  much  more 
likely  to  possess  the  proper  affinities  for  uniting  with  the  tissues 
of  the  same  sex,  and  thus  becoming  developed,  than  witli  those  of 
the  opposite  sex. 

I  was  first  led  to  infer  that  a  relation  of  this  kind  exists,  from 
the  fact  that  whenever  and  in  whatever  manner  the  adult  male 
differs  from  the  adult  female,  he  differs  in  the  same  manner  from 
the  young  of  both  sexes.  The  generality  of  this  fact  is  quite  re- 
markable: it  holds  good  with  almost  all  mammals,  birds,  amphib- 
ians, and  fishes;  also  with  many  crustaceans,  spiders,  and  some 
few  insects,  such  as  certain  orthoptera  and  libellulae.    In  all  these 

"should  have  sug-gested  the  possibility  of  modifying'  the  sexual  colors 
"of  birds  by  a  course  of  artificial  selection.  When  he  did  so,  he  was 
"in  ignorance  of  these  facts  that  I  have  related;  but  it  is  remarkable 
"how  very  closely  he  suggested  the  right  method  of  procedure." 

ss  References  are  given  in  my  'Variation  of  Animals  under  Domes- 
tication,' vol.  ii.  p.  72. 


22g  THE   DESCENT  OF  MAN. 

cases  the  variations,  through  the  accumulation  of  which  the  male 
acquired  his  proper  masculine  characters,  must  have  occurred  at  a 
somewhat  late  period  of  life;  otherwise  the  young  males  would 
have  been  similarly  characterized;  and  comformahly  with  our  rule, 
the  variations  are  transmitted  to  and  developed  in  the  adult  males 
alone.  When,  on  the  other  hand,  the  adult  male  closely  resembles 
the  young  of  both  sexes  (these,  with  rare  exceptions,  being  alike), 
he  generally  resembles  the  adult  female;  and  in  most  of  these  cases 
the  variations  through  which  the  young  and  old  acquired  their 
present  characters,  probably  occurred,  according  to  our  rule,  dur- 
ing youth.  But  there  is  here  room  for  doubt,  for  characters  are 
sometimes  transferred  to  the  offspring  at  an  earlier  age  than  that 
at  which  they  first  appeared  in  the  parents,  so  that  the  parents 
may  have  varied  when  adult,  and  have  transferred  their  characters 
to  their  offspring  whilst  young.  There  are,  moreover,  many  ani- 
mals, in  which  the  two  sexes  closely  resemble  each  other,  and  yet 
both  differ  from  their  young;  and  here  the  characters  of  the  adults 
must  have  been  acquired  late  in  life;  nevertheless,  these  char- 
acters, in  apparent  contradiction  to  our  rule,  are  transferred  to 
both  sexes.  We  must  not,  however,  overlook  the  possibility  or 
even  probability  of  successive  variations  of  the  same  nature  occur- 
ring, under  exposure  to  similar  conditions,  simultaneously  in  both 
sexes  at  a  rather  late  period  of  life;  and  in  this  case  the  variations 
would  be  transferred  to  the  offspring  of  both  sexes  at  a  corre- 
sponding late  age;  and  there  would  then  be  no  real  contradiction 
to  the  rule  that  variations  occurring  late  in  life  are  transferred 
exclusively  to  the  sex  in  which  they  first  appeared.  This  latter 
rule  seems  to  hold  true  more  generally  than  the  second  one, 
namely,  that  variations  which  occur  in  either  sex  early  in  life  tend 
to  be  transferred  to  both  sexes.  As  it  was  obviouslj^  impossible 
even  to  estimate  in  how  large  a  number  of  cases  throughout  the 
animal  kingdom  these  two  propositions  held  good,  it  occurred  to 
me  to  investigate  some  striking  or  crucial  instances,  and  to  rely 
on  the  result. 

An  excellent  case  for  investigation  is  afforded  by  the  Deer 
family.  In  all  the  species,  but  one,  the  horns  are  developed  only 
in  the  males,  though  certainly  transmitted  through  the  females, 
and  capable  of  abnormal  development  in  them.  In  the  reindeer, 
on  the  other  hand,  the  female  is  provided  with  horns;  so  that  in 
this  species,  the  horns  ought,  according  to  our  rule,  to  appear  early 
in  life,  long  before  the  two  sexes  are  mature  and  have  come  to  dif- 
fer much  in  constitution.  In  all  the  other  species  the  horns  ought 
to  appear  later  in  life,  which  would  lead  to  their  development  in 
that  sex  alone,  in  which  they  first  appeared  in  the  progenitor  of 
the  whole  Family.  Now  in  seven  species,  belonging  to  distinct 
sections  of  the  family  and  inhabiting  different  regions,  in  which 
the  stags  alone  bear  horns,  I  find  that  the  horns  first  appear  at 


•  SEXUAL   SELECTION.  229 

periods,  varying  from  nine  monttis  after  birth  in  tlie  roebuck,  to 
ten,  twelve  or  even  more  months  in  the  stags  of  the  six  other 
and  larger  species.^^  But  with  the  reindeer  the  case  is  widely  dif- 
ferent; for,  as  I  hear  from  Prof.  Nilsson,  who  kindly  made  special 
inquiries  for  me  in  Lapland,  the  horns  appear  in  the  young  ani- 
mals within  four  or  five  weeks  after  birth,  and  at  the  same  time 
in  both  sexes.  So  that  here  we  have  a  structure,  developed  at  a 
most  unusually  early  age  in  one  species  of  the  family,  and  like- 
wise common  to  both  sexes  in  this  one  species  alone. 

In  several  kinds  of  antelopes,  only  the  males  are  provided  with 
horns,  whilst  in  the  greater  number  both  sexes  bear  horns.  With 
respect  to  the  period  of  development,  Mr.  Blyth  informs  me  that 
there  was  at  one  time  in  the  Zoological  Gardens  a  young  koodoo 
(Ant.  strepsiceros),  of  which  the  males  alone  are  horned,  and  also 
the  young  of  a  closely-allied  species,  the  eland  (Ant.  oreas),  in 
which  both  sexes  are  horned.  Now  it  is  in  strict  conformity  with 
our  rule,  that  in  the  young  male  koodoo,  although  ten  months  old, 
the  horns  were  remarkably  small,  considering  the  size  ultimately 
attained  by  them;  whilst  in  the  young  male  eland,  although  only 
three  months  old,  the  horns  were  already  very  much  larger  than 
in  the  koodoo.  It  is  also  a  noticeable  fact  that  in  the  prong-horned 
antelope,*"  only  a  few  of  the  females,  about  one  in  five,  have  horns 
and  these  are  in  a  rudimentary  state,  though  sometimes  above  four 
inches  long;  so  that  as  far  as  concerns  the  possession  of  horns  by 
the  males  alone,  this  species  is  in  an  intermediate  condition,  and 
the  horns  do  not  appear  until  about  five  or  six  months  after  birth. 
Therefore  in  comparison  with  what  little  we  know  of  the  develop- 
ment of  the  horns  in  other  antelopes,  and  from  what  we  do  know 
with  respect  to  the  horns  of  deer,  cattle,  &c.,  those  of  the  prong- 
horned  antelope  appear  at  an  intermediate  period  of  life,— that  is, 
not  very  early,  as  in  cattle  and  sheep,  nor  very  late,  as  in  the  larg- 
er deer  and  antelopes.  The  horns  of  sheep,  goats,  and  cattle, 
which  are  well  developed  in  both  sexes,  though  not  quite  equal  in 


^^  I  am  much  obliged  to  Mr.  Cupples  for  having  made  inquiries 
for  me  in  regard  to  the  Roebuck  and  Red  Deer  of  Scotland  from  Mr. 
Robertson,  the  experienced  head-forester  to  the  Marquis  of  Breadal- 
bane.  In  regard  to  Fallow-deer,  I  have  to  thank  Mr.  Eyton  and 
others  for  information.  For  the  Cervus  aloes  of  N.  America,  see 
'Land  and  Water,'  1868,  pp.  221  and  254;  and  for  the  C.  Virginianus 
and  strongyliceros  of  the  same  continent,  see  J.  D.  Caton,  in  'Ottawa 
Acad,  of  Nat.  Sc.  1868,  p.  13.  For  Cervus  Eldi  of  Pegu,  see  Lieut. 
Beavan,    'Proc.   Zoolog.    Soc*   1867,   p.   762. 

^Antilocapra  Americana.  I  have  to  thank  Dr.  Canfield  for  In- 
formation with  respect  to  the  horns  of  the  female:  see  also  his  paper 
in  'Proc.  Zoolog.  Soc'  1866,  p.  109.  Also  Owen.  'Anatomy  of  Verte- 
brates,' vol.  iii.  p.  627. 

16 


230  THE  DESCENT  OP  MAN. 

size,  can  be  felt,  or  even  seen,  at  birth  or  soon  afterwards.'^  Our 
rule,  however,  seems  to  fail  in  some  breeds  of  sheep,  for  instance 
merinos,  in  which  the  rams  alohe  are  horned;  for  I  cannot  find 
on  inquiry,*^  that  the  horns  are  developed  later  in  life  in  this 
breed  than  in  ordinary  sheep  in  which  both  sexes  are  horned.  But 
with  domesticated  sheep  the  presence  or  absence  of  horns  is  not  a 
firmly  fixed  character;  for  a  certain  proportion  of  the  merino  ewes 
bear  small  horns,  and  some  of  the  rams  are  hornless;  and  in  most 
breeds  hornless  ewes  are  occasionally  produced. 

Dr.  W.  Marshall  has  lately  made  a  special  study  of  the  protuber- 
ances so  common  on  the  heads  of  birds,*^  and  he  comes  to  the  fol- 
lowing conclusion; — that  with  those  species  in  which  they  are  con- 
fined to  the  males,  they  are  developed  late  in  life;  whereas  with 
those  species  in  which  they  are  common  to  the  two  sexes,  they  are 
developed  at  a  very  early  period.  This  is  certainly  a  strildng  con- 
firmation of  my  two  laws  of  inheritance. 

In  most  of  the  species  of  the  splendid  family  of  the  Pheasants, 
the  males  differ  conspicuously  from  the  females,  and  they  acquire 
their  ornaments  at  a  rather  late  period  of  life.  The  eared  pheas- 
ant (Crossoptilon  auritum),  however,  offers  a  remarkable  excep- 
tion, for  both  sexes  possess  the  fine  cau'''al  plumes,  the  large  ear- 
tufts  and  the  crimson  velvet  about  the  head;  I  find  that  all  these 
characters  appear  very  early  in  life  in  accordance  with  rule.  The 
adult  male  can,  however,  be  distinguished  from  the  adult  female 
by  the  presence  of  spurs;  and  conformably  with  our  rule,  these  do 
not  begin  to  be  developed  before  the  age  of  six  months,  as  I  am 
assured  by  Mr.  Bartlett,  and  even  at  this  age,  the  two  sexes  can 
hardly  be  distinguished.^     The  male  and  female  Peacock  differ 


*i  I  have  been  assured  that  the  horns  of  the  sheep  in  North  Wales 
can  always  be  felt,  and  are  sometimes  even  an  inch  in  length,  at 
birth.  Youatt  says  ('Cattle,'  1834,  p.  277),  that  the  prominence  of  the 
frontal  bone  in  cattle  penetrates  the  cutis  at  birth,  and  that  the 
horny  matter  is  soon  formed  over  it. 

^  I  am  greatly  indebted  to  Prof.  Victor  Carus  for  having  made 
inquiries  for  me,  from  the  highest  authorities,  with  respect  to  the 
merino  sheep  of  Saxony.  On  the  Guinea  coast  of  Africa  there  is, 
however,  a  breed  of  sheep  in  which,  as  with  merinos,  the  rams  alone 
bear  horns;  and  Mr.  Winwood  Reade  informs  me  that  in  one  case 
observed  by  him,  a  young  ram,  born  on  Feb.  10th,  first  showed  horns 
on  March  6th,  so  that  in  this  instance,  in  conformity  with  rule,  the 
development  of  the  horns  occurred  at  a  later  period  of  life  than  in 
Welsh  sheep,   in  which  both  sexes  are  horned, 

"  'Ueber  die  knochernen  Schadelhocker  der  Vogel'  in  the  'Nieder- 
landischen  Archiv  fur  Zoologie,'  Band  I.  Heft  2,  1872. 

**  In  the  common  peacock  (Pavo  cristatus)  the  male  alone  possesses 
spurs,  whilst  both  sexes  of  the  Java  Peacock  (P.  muticus)  offer  the 
unusual  case  of  being  furnished  with  spurs.  Hence  I  fully  expected 
that  in  the  latter  species  they  would  have  been  developed  earlier  ic 


SEXUAL    SELECTION.  231 

conspicuously  from  each  other  in  almost  every  part  of  their  plum- 
age, except  in  the  elegant  head-crest,  which  is  common  to  both 
sexes;  and  this  is  developed  very  early  in  life,  long  before  the  other 
ornaments,  which  are  confined  to  the  male.  The  wild-duck  offers 
an  analogous  case,  for  the  beautiful  green  speculum  on  the  wings 
is  common  to  both  sexes,  though  duller  and  somewhat  smaller  in 
the  female,  and  it  is  developed  early  in  life,  v/hilst  the  curled  tail- 
feathers  and  other  ornaments  of  the  male  are  developed  later. *^ 
Between  such  extreme  cases  of  close  sexual  resemblance  and  wide 
dissimilarity,  as  those  of  the  Crossoptilon  and  peacock,  many  in- 
termediate ones  could  be  given,  in  which  the  characters  follow  our 
two  rules  in  their  order  of  development. 

As  most  insects  emerge  from  the  pupal  state  in  a  mature  con- 
dition, it  is  doubtful  whether  the  period  of  development  can  deter- 
mine the  transference  of  their  characters  to  one  or  to  both  sexes. 
But  we  do  not  know  that  the  colored  scales,  for  instance,  in  two 
species  of  butterflies,  in  one  of  which  the  sexes  differ  in  color, 
whilst  in  the  other  they  are  alike,  are  developed  at  the  same  rela- 
tive age  in  the  cocoon.  Nor  do  we  know  whether  all  the  scales 
are  simultaneously  developed  on  the  wings  of  the  same  species 
of  butterfly,  in  which  certain  colored  marks  are  confined  to  one 
sex,  whilst  others  are  common  to  both  sexes.  A  difference  of  this 
kind  in  the  period  of  development  is  not  so  improbable  as  it  may 
at  first  appear;  for  with  the  Orthoptera,  which  assume  their 
adult  state,  not  by  a  single  metamorphosis,  but  by  a  succession 
of  moults,  the  young  males  of  some  species  at  first  resemble  the 
females,  and  acquire  their  distinctive  masculine  characters  only 


life  than  in  the  common  peacock;  but  M.  Hogt  of  Amsterdam  in- 
forms me,  that  with  young  birds  of  the  previous  year,  of  both  species, 
compared  on  April  23rd,  1869,  there  was  no  difference  in  the  develop- 
ment of  the  spurs.  The  spurs,  however,  were  as  yet  represented 
merely  by  slight  knobs  or  elevations.  I  presume  that  I  should  have 
been  informed  if  any  difference  in  the  rate  of  development  had  been 
observed  subsequently. 

*s  In  some  other  species  of  the  Duck  family  the  speculum  differs 
In  a  greater  degree  in  the  two  sexes;  but  I  have  not  been  able  to 
discover  whether  its  full  development  occurs  later  in  life  in  the 
males  of  such  species,  than  in  the  male  of  the  common  duck,  as 
ought  to  be  the  case  according  to  our  rule.  With  the  allied  Mergus 
cucullatus  we  have,  however,  a  case  of  this  kind:  the  two  sexes  dif- 
fer conspicuously  in  general  plumage,  and  to  a  considerable  degree 
In  the  speculum,  which  is  pure  white  in  the  male  and  grayish-white 
in  the  female.  Now  the  young  males  at  first  entirely  resemble  the 
females,  and  have  a  grayish-white  speculum,  which  becomes  pure 
white  at  an  earlier  age  than  that  at  which  the  adult  male  acquires 
his  other  and  more  strongly-marked  sexual  differences:  see  Audu« 
bon,   'Ornithological  Biography,'  vol.  iii.   1835,  pp.  249-250. 


232  THE  DESCENT  OF  MAN. 

at  a  later  moult.  Strictly  analogous  cases  occur  at  the  successive 
moults  of  certain  male  crustaceans. 

We  have  as  yet  considered  the  transference  of  characters,  rela- 
tively to  their  period  of  development,  only  in  species  in  a  natural 
state;  we  will  now  turn  to  domesticated  animals,  and  first  toucn 
on  monstrosities  and  diseases.  The  presence  of  supernumerary 
digits,  and  the  absence  of  certain  phalanges,  must  be  determined 
at  an  early  embryonic  period — the  tendency  to  profuse  bleeding  is 
at  least  congenital,  as  is  probably  color-blindness — yet  these  pe- 
culiarities, and  other  similar  ones,  are  often  limited  in  their  trans- 
mission to  one  sex;  so  that  the  rule  that  characters,  developed  at 
an  early  period,  tend  to  be  transmitted  to  both  sexes,  here  wholly 
fails.  But  this  rule,  as  before  remarked,  does  not  appear  to  be 
nearly  so  general  as  the  converse  one,  namely,  that  characters 
which  appear  late  in  life  in  one  sex  are  transmitted  exclusively  to 
the  same  sex.  From  the  fact  of  the  above  abnormal  peculiarities 
becoming  attached  to  one  sex,  long  before  the  sexual  functions  are 
active,  we  may  infer  that  there  must  be  some  difference  between 
the  sexes  at  an  extremely  early  age.  With  respect  to  sexually  lim- 
ited diseases,  we  know  too  little  of  the  period  at  which  they  orig- 
inate, to  draw  any  safe  conclusion.  Gout,  however,  seems  to  fall 
under  our  rule,  for  it  is  generally  caused  by  intemperance  during 
manhood,  and  is  transmitted  from  the  father  to  his  sons  in  a 
much  more  marked  manner  than  to  his  daughters. 

In  the  various  domestic  breeds  of  sheep,  goats,  and  cattle,  the 
males  differ  from  their  respective  females  in  the  shape  or  develop- 
ment of  their  horns,  forehead,  mane,  dewlap,  tail,  and  hump  on 
the  shoulders;  and  these  peculiarities,  in  accordance  with  our 
rule,  are  not  fully  developed  antil  a  rather  late  period  of  life. 
The  sexes  of  dogs  do  not  differ,  except  that  in  certain  breeds, 
especially  in  the  Scotch  deer-hound,  the  male  is  much  larger 
and  heavier  than  the  female;  and,  as  we  shall  see  in  a  future 
chapter,  the  male  goes  on  increasing  in  size  to  an  unusually  late 
period  of  life,  which,  according  to  rule,  will  account  for  his  in- 
creased size  being  transmitted  to  his  male  offspring  alone.  On 
the  other  hand,  the  tortoise-shell  color,  which  is  confined  to 
female  cats,  is  quite  distinct  at  birth,  and  this  case  violates  the 
rule.  There  is  a  breed  of  pigeons  in  which  the  males  alone  are 
streaked  with  black,  and  the  streaks  can  be  detected  even  in  the 
nestlings;  but  they  become  more  conspicuous  at  each  successive 
moult,  so  that  this  case  partly  opposes  and  partly  supports  the 
rule.  With  the  English  Carrier  and  Pouter  pigeons,  the  full  de- 
velopment of  the  wattle  and  the  crop  occurs  rather  late  in  life, 
and  conformably  with  the  rule,  these  characters  are  transmitted 
in  full  perfection  to  the  males  alone.  The  following  cases  perhaps 
come  within  the  class  previously  alluded  to,  in  which  both  sexes 
have  varied  in  the  same  manner  at  a  rather  late  period  of  life, 


SEXUAL    SELECTION.  233 

md  have  consequently  transferred  their  new  characters  to  both 
sexes  at  a  corresponding  late  period;  and  if  so,  these  cases  are 
not  opposed  to  our  rule: — there  exist  sub-breeds  of  the  pigeon, 
described  by  Neumeister,*^  in  which  both  sexes  change  their  color 
during  two  or  three  moults  (as  is  likewise  the  case  with  the 
Almond  Tumbler),  nevertheless,  these  changes,  though  occurring 
rather  late  in  life,  are  common  to  both  sexes.  One  variety  of  the 
Canary-bird,  namely  the  London  Prize,  offers  a  nearly  analogous 
case. 

With  the  breeds  of  the  Fowl  the  inheritance  of  various  charac- 
ters by  one  or  both  sexes,  seems  generally  determined  by  the 
period  at  which  such  characters  are  developed.  Thus  in  all  the 
many  breeds  in  which  the  adult  male  differs  greatly  in  color  from 
the  female,  as  well  as  from  the  wild  parent-species,  he  differs 
also  from  the  young  male,  so  that  the  newly-acquired  characters 
must  have  appeared  at  a  rather  late  period  of  life.  On  the  other 
hand,  in  most  of  the  breeds  in  which  the  two  sexes  resemble  each 
other,  the  young  are  colored  in  nearly  the  same  manner  as  their 
parents,  and  this  renders  it  probable  that  their  colors  first  ap- 
peared early  in  life.  We  have  instances  of  this  fact  in  all  black 
and  white  breeds,  in  which  the  young  and  old  of  both  sexes  are 
alike;  nor  can  it  be  maintained  that  there  is  something  peculiar 
in  a  black  or  white  plumage,  which  leads  to  its  transference  to 
both  sexes;  for  the  males  alone  of  many  natural  species  are  either 
black  or  white,  the  females  being  differently  colored.  With  the 
so-called  Cuckoo  sub-breeds  of  the  fowl,  in  which  the  feathers  are 
transversely  pencilled  with  dark  stripes,  both  sexes  and  the  chick- 
ens are  colored  in  nearly  the  same  manner.  The  laced  plumage 
of  the  Sebright  bantam  is  the  same  in  both  sexes,  and  in  the  young 
chickens  the  wing-feathers  are  distinctly,  though  imperfectly 
laced.  Spangled  Hamburgs,  however,  offer  a  partial  exception; 
for  the  two  sexes,  though  not  quite  alike,  resemble  each  other 
more  closely  than  do  the  sexes  of  the  aboriginal  parent-species; 
yet  they  acquire  their  characteristic  plumage  late  in  life,  for  the 
chickens  are  distinctly  pencilled.  With  respect  to  other  characters 
besides  color,  in  the  wild-parent  species  and  in  most  of  the  domes- 
tic breeds,  the  males  alone  possess  a  well-developed  comb;  but  in 
the  young  of  the  Spanish  fowl  it  is  largely  developed  at  a  very 
early  age,  and,  in  accordance  with  this  early  development  in  the 
male,  it  is  of  unusual  size  in  the  adult  female.  In  the  Game 
breeds,  pugnacity  is  developed  at  a  wonderfully  early  age,  of 
which  curious  proofs  could  be  given;  and  this  character  is  trans- 
mitted to  both  sexes,  so  that  the  hens,  from  their  extreme  pug- 

*8  'Das  Ganze  der  Taubenzucht,'  1837,  s.  21,  24.  For  the  case  of  the 
streaked  pigeons,  see  Dr.  Chapuis,  'Le  pigeon  voyageur  Beige,'  1865, 
p.  87. 


234  THE  DESCENT  OF  MAN. 

nacity,  are  now  generally  exhibited  in  separate  pens.  With  the 
Polish  breeds  the  bony  protuberance  of  the  skull  which  supports 
the  crest  is  partially  developed  even  before  the  chickens  are 
hatched,  and  the  crest  itself  soon  begins  to  grow,  though  at  first 
feebly  ;^^  and  in  this  breed  the  adults  of  both  sexes  are  charac- 
terized by  a  great  bony  protuberance  and  an  immense  crest. 

Finally,  from  what  we  have  now  seen  of  the  relation  which 
exists  in  many  natural  species  and  domesticated  races,  between 
the  period  of  the  development  of  their  characters  and  the  manner 
of  their  transmission — for  example,  the  striking  fact  of  the  early 
grov/th  of  the  horns  in  the  reindeer,  in  which  both  sexes  bear 
horns,  in  comparison  with  their  much  later  growth  in  the  other 
species  in  which  the  male  alone  bears  horns — we  may  conclude 
that  one,  though  not  the  sole  cause  of  characters  being  exclusively 
inherited  by  one  sex,  is  their  development  at  a  late  age.  And 
secondly  that  one,  though  apparently  a  less  efiicient  cause  of  char- 
acters being  inherited  by  both  sexes,  is  their  development  at  an 
early  age,  whilst  the  sexes  differ  but  little  in  constitution.  It 
appears,  however,  that  some  difference  must  exist  between  the 
sexes  even  during  a  very  early  embryonic  period,  for  characters 
developed  at  this  age  not  rarely  become  attached  to  one  sex. 

Summary  and  concluding  remarks. — From  the  foregoing  dis- 
cussion on  the  various  laws  of  inheritance,  we  learn  that  the 
characters  of  the  parents  often,  or  even  generally,  tend  to  become 
developed  in  the  offspring  of  the  same  sex,  at  the  same  age,  and 
periodically  at  the  same  season  of  the  year,  in  which  they  first 
appeared  in  the  parents.  But  these  rules,  owing  to  unknown 
causes,  are  far  from  being  fixed.  Hence  during  the  modification 
of  a  species,  the  successive  changes  may  readily  be  transmitted 
in  different  ways;  some  to  one  sex,  and  some  to  both;  some  to 
the  offspring  at  one  age,  and  some  to  the  offspring  at  all  ages. 
Not  only  are  the  laws  of  inheritance  extremely  complex,  but  so 
are  the  causes  which  induce  and  govern  variability.  The  varia- 
tions thus  induced  are  preserved  and  accumulated  by  sexual  selec- 
tion, which  is  in  itself  an  extremely  complex  affair,  depending,  as 
it  does,  on  the  ardor  in  love,  the  courage,  and  the  rivalry  of  the 
males,  as  well  as  on  the  powers  of  perception,  the  taste,  and  will 
of  the  female.  Sexual  selection  will  also  be  largely  dominated  by 
natural  selection  tending  towards  the  general  welfare  of  the  spe- 
cies.   Hence  the  manner  in  which  the  individuals  of  either  or  both 


"  For  full  particulars  and  references  on  all  these  points  respecting- 
the  several  breeds  of  the  Fowl,  see  'Variation  of  Animals  and  Plants 
under  Domestication,'  vol.  i.  pp.  250,  256.  In  regard  to  the  higher  ani- 
mals, the  sexual  differences  v/hich  have  arisen  under  domestication 
are  described  in  the  same  work  under  the  head  of  each  species. 


SEXUAL    SELECTION.  235 

sexes  have  been  affected  through  sexual  selection  cannot  fail  to 
be  complex  in  the  highest  degree. 

When  variations  occur  late  in  life  in  one  sex,  and  are  trans- 
mitted to  the  same  sex  at  the  same  age,  the  other  sex  and  the 
young  are  left  unmodified.  When  they  occur  late  in  life,  but 
are  transmitted  to  both  sexes  at  the  same  age,  the  young  alone 
are  left  unmodified.  Variations,  however,  may  occur  at  any  period 
of  life  in  one  sex  or  in  both,  and  be  transmitted  to  both  sexes 
at  all  ages,  and  then  all  the  individuals  of  the  species  are  sim- 
ilarly modified.  In  the  following  chapters  it  will  be  seen  that  all 
these  cases  frequently  occur  in  nature. 

Sexual  selection  can  never  act  on  any  animal  before  the  age 
for  reproduction  arrives.  From  the  great  eagerness  of  the  "male 
it  has  generally  acted  on  this  sex  and  not  on  the  females.  The 
males  have  thus  become  provided  with  weapons  for  fighting  with 
their  rivals,  with  organs  for  discovering  and  securely  holding  the 
female,  and  for  exciting  or  charming  her.  V/hen  the  sexes  differ 
in  these  respects,  it  is  also,  as  we  have  seen,  an  extremely  general 
law  that  the  adult  male  differs  more  or  less  from  the  young  male; 
and  we  may  conclude  from  this  fact  that  the  successive  variations, 
by  which  the  adult  male  became  modified,  did  not  generally  occur 
much  before  the  age  for  reproduction.  Whenever  some  or  many 
of  the  variations  occurred  early  in  life,  the  young  males  would  par- 
take more  or  less  of  the  characters  of  the  adult  males;  and  dif- 
ferences of  this  kind  between  the  old  and  young  males  may  be 
observed  in  many  species  of  animals. 

It  is  probable  that  young  male  animals  have  often  tended  to 
vary  in  a  manner  which  would  not  only  have  been  of  no  use  to 
them  at  an  early  age,  but  would  have  been  actually  injurious — 
as  by  acquiring  bright  colors,  which  would  render  them  con- 
spicuous to  their  enemies,  or  by  acquiring  structures,  such  as 
great  horns,  which  would  expend  much  vital  force  in  their  devel- 
opment. Variations  of  this  kind  occurring  in  the  young  males 
would  almost  certainly  be  eliminated  through  natural  selection. 
With  the  adult  and  experienced  males,  on  the  other  hand,  the 
advantages  derived  from  the  acquisition  of  such  characters,  would 
more  than  counterbalance  some  exposure  to  danger,  and  some 
loss  of  vital  force. 

As  variations  which  give  to  the  male  a  better  chance  of  con- 
quering other  males,  or  of  finding,  securing,  or  charming  the  oppo- 
site sex,  would,  if  they  happen  to  arise  in  the  female,  be  of  no 
service  to  her,  they  would  not  be  preserved  in  her  through  sexual 
selection.  We  have  also  good  evidence  with  domesticated  ani- 
mals, that  variations  of  all  kinds  are,  if  not  carefully  selected, 
soon  lost  through  intercrossing  and  accidental  deaths.  Conse- 
quently in  a  state  of  nature,  if  variations  of  the  above  kind 
chanced  to  arise  in  the  female  line,  and  to  be  transmitted  exclu- 


236  THE  DESCENT  OF  MAN. 

sively  in  this  line,  they  would  be  extremely  liable  to  be  lost.  If, 
however,  the  females  varied  and  submitted  their  newly  acquired 
characters  to  their  offsprings  of  both  sexes,  the  characters  which 
were  advantageous  to  the  males  would  be  preserved  by  them 
through  sexual  selection,  and  the  two  sexes  would  in  consequence 
be  modified  in  the  same  manner,  although  such  characters  were 
of  no  use  to  the  females;  but  I  shall  hereafter  have  to  recur  to 
these  more  intricate  contingencies.  Lastly,  the  females  may  ac- 
quire, and  apparently  have  often  acquired  by  transference,  char- 
acters from  the  male  sex. 

As  variations  occurring  late  in  life,  and  transmitted  to  one  sex 
alone,  have  incessantly  been  taken  advantage  of  and  accumulated 
through  sexual  selection  in  relation  to  the  reproduction  of  the 
species;  therefore  it  appears,  at  first  sight,  an  unaccountable  fact 
that  similar  variations  have  not  frequently  been  accumulated 
through  natural  selection,  in  relation  to  the  ordinary  habits  of  life. 
If  this  had  occurred,  the  two  sexes  would  often  have  been  differ- 
ently modified,  for  the  sake,  for  instance,  of  capturing  prey  or  of 
escaping  from  danger.  Differences  of  this  kind  between  the  two 
sexes  do  occasionally  occur,  especially  in  the  lower  classes.  But 
this  implies  that  the  two  sexes  follow  different  habits  in  their 
struggles  for  existence,  which  is  a  rare  circumstance  with  the 
higher  animals.  The  case,  however,  is  widely  different  with  the 
reproductive  functions,  in  which  respect  the  sexes  necessarily  dif- 
fer. For  variations  in  structure  which  are  related  to  these  func- 
tions, have  often  proved  of  value  to  one  sex,  and  from  having 
arisen  at  a  late  period  of  life,  have  been  transmitted  to  one  sex 
alone;  and  such  variations,  thus  preserved  and  transmitted, 
have  given  rise  to  secondary  sexual  characters. 

In  the  following  chapters,  I  shall  treat  of  the  secondary  sexual 
characters  in  animals  of  all  classes,  and  shall  endeavor  in  each 
case  to  apply  the  principles  explained  in  the  present  chapter.  The 
lowest  classes  will  detain  us  for  a  very  short  time,  but  the  higher 
animals,  especially  birds,  must  be  treated  at  considerable  length. 
It  should  be  borne  in  mind  that  for  reasons  already  assigned,  I 
intend  to  give  only  a  few  illustrative  instances  of  the  innumerable 
structures  by  the  aid  of  which  the  male  finds  the  female,  or  when 
found,  holds  her.  On  the  other  hand,  all  structures  and  instincts 
by  the  aid  of  which  the  male  conquers  other  males,  and  by  whicn 
he  allures  or  excites  the  female,  will  be  fully  discussed,  as  these 
are  in  many  ways  the  most  interesting. 


Supplement  on  the  proportional  numbers  of  the  two  sexes  in  an- 
imals belonging  to  various  classes 

As  no  one,  as  far  as  I  can  discover,  has  paid  attention  to  the 
relative  numbers  of  the  two  sexes  throughout  the  animal  king- 


PROPORTION   OF    THE    SEXES.  237 

dom,  I  will  here  give  such  materials  as  I  have  been  able  to  collect, 
althougli  tliey  are  extremely  imperfect.  They  consist  in  only  a 
few  instances  of  actual  enumeration,  and  the  numbers  are  not 
very  large.  As  the  proportions  are  known  with  certainty  only  in 
mankind,  I  will  first  give  them  as  a  standard  of  comparison. 

Man. — In  England  during  ten  years  (from  1857  to  1866)  the 
average  number  of  children  born  alive  yearly  was  707,120,  in 
the  proportion  of  104.5  males  to  100  females.  But  in  1857  the 
male  births  throughout  England  were  as  105.2  and  in  1865  as 
104.0  to  100.  Looking  to  separate  districts,  in  Buckinghamshire 
(where  about  5000  children  are  annually  born)  the  mean  propor- 
tion of  male  to  female  births,  during  the  whole  period  of  the  above 
ten  years  was  as  102.8  to  100;  whilst  in  N.  Wales  (where  the 
average  annual  births  are  12,873)  it  was  as  high  as  106.2  to  100. 
Taking  a  still  smaller  district,  viz.,  Rutlandshire  (where  the 
annual  births  average  only  739),  in  1864  the  male  births  were  as 
114.6,  and  in  1862  as  only  97.0  to  100;  but  even  in  this  small  dis- 
trict the  average  of  the  7385  births  during  the  whole  ten  years, 
was  as  104.5  to  100;  that  is  in  the  same  ratio  as  throughout  Eng- 
land.^ The  proportions  are  sometimes  slightly  disturbed  by  un- 
known causes;  thus  Prof.  Faye  states  "that  in  some  districts  of 
"Norway  there  has  been  during  a  decennial  period  a  steady  de- 
"ficiency  of  boys,  whilst  in  others  the  opposite  condition  has 
"existed."  In  Prance  during  forty-four  years  the  male  to  the 
female  births  have  been  as  106.2  to  100;  but  during  this  period 
it  has  occurred  five  times  in  one  department,  and  six  times  in. 
another,  that  the  female  births  have  exceeded  the  males.  In 
Russia  the  average  proportion  is  as  high  as  108.9,  and  in  Phila- 
delphia in  the  United  States  as  110.5  to  100."^  The  average  for 
Europe,  deduced  by  Bickes  from  about  seventy  million  births, 
is  106  males  to  100  females.  On  the  other  hand,  with  white  chil- 
dren born  at  the  Cape  of  Good  Hope,  the  proportion  of  males  is  so 
low  as  to  fluctuate  during  successive  years  between  90  and  99 
males  for  every  100  females.  It  is  a  singular  fact  that  with  Jews 
the  proportion  of  male  births  is  decidedly  larger  than  with  Chris- 
tians: thus  in  Prussia  the  proportion  is  as  113,  in  Breslau  as 
114,  and  in  Livonia  as  120  to  100;    the  Christian  births  in  these 

*8  'Twenty-ninth  Annual  Report  of  the  Registrar-General  for  1866.' 
In  this  report  (p.  xii.)  a  special  decennial  table  is  given. 

*9  For  Norway  and  Russia,  see  abstract  of  Prof.  Faye's  reseai*ches, 
in  'British  and  Foreign  Medico-Chirurg.  Review,'  April,  1867,  pp.  343, 
345.  For  France,  the  'Annuaire  pour  I'An  1867,'  p.  213.  For  Philadel- 
phia, Dr.  Stockton-Hough,  'Social  Science  Assoc'  1874.  For  the  Cape 
of  Good  Hope,  Quetelet  as  quoted  by  Dr.  H.  H.  Zouteveen,  in  the 
Dutch  Translation  of  this  work  (vol.  i.  p.  417),  where  much  informa- 
tion is  given  on  the  proportion  of  the  sexes. 


238  THE  DESCENT  OF  MAN. 

countries  being  the  same  as  usual,  for  instance,  in  Livonia  as  104 
to  100.^° 

Prof.  Faye  remarks  that  "a  still  greater  preponderance  of  males 
"would  be  met  with,  if  death  struck  both  sexes  in  equal  pro- 
"portion  in  the  womb  and  during  birth.  But  the  fact  is,  that  for 
"every  100  still-born  females,  we  have  in  several  countries  from 
"134.6  to  144.9  still-born  males.  During  the  first  four  or  five  years 
"of  life,  also,  more  male  children  die  than  females;  for  example 
"in  England,  during  the  first  year,  126  boys  die  for  every  100 
"girls— a  proportion  which  in  France  is  still  more  unfavorable."" 
Dr.  Stockton-Hough  accounts  for  these  facts  in  part  by  the  more 
frequent  defective  development  of  males  than  of  females.  We 
have  before  seen  that  the  male  sex  is  more  variable  iii  structure 
than  the  female;  and  variations  in  important  organs  would  gen- 
erally be  injurious.  But  the  size  of  the  body,  and  especially  of  the 
head,  being  greater  in  male  than  female  infants  is  another  cause; 
for  the  males  are  thus  more  liable  to  be  injured  during  parturition. 
Consequently  the  still-born  males  are  more  numerous;  and,  as  a 
highly  competent  judge.  Dr.  Crichton  Browne,'^  believes,  male  in- 
fants often  suffer  in  health  for  some  years  after  birth.  Owing  to 
this  excess  in  the  death-rate  of  male  children,  both  at  birth  and 
for  some  time  subsequently,  and  owing  to  the  exposure  of  grown 
men  to  various  dangers,  and  to  their  tendency  to  emigrate,  the 
females  in  all  old-settled  countries,  where  statistical  records  have 
been  kept,^^  are  found  to  preponderate  considerably  over  the  males. 

It  seems  at  first  sight  a  mysterious  fact  that  in  different  na- 
tions, under  different  conditions  and  climates,  in  Naples,  Prussia, 
Westphalia,  Holland,  France,  England  and  the  United  States,  the 

^0  In  regard  to  the  Jews,  see  M.  Tliury,  'La  Loi  de  Production  des 
Sexes,'  1863,  p.  25. 

51  'British  and  Foreign  Medico-Chirurg.  Review,'  April,  1867,  p.  343. 
Dr.  Stark  also  remarks  ('Tenth  Annual  Report  of  Births,  Deaths, 
&c.,  in  Scotland,'  1867,  p.  xxviii.)  that  "These  examples  maj^  suffice 
"to  show  that,  at  almost  every  stage  of  life,  the  males  in  ,Scotland 
"have  a  greater  liability  to  death  and  a  higher  death-rate  than  the 
"females.  The  fact,  however,  of  this  peculiarity  being  most  strongly 
"developed  at  that  infantile  period  of  life  when  the  dress,  food,  and 
"general  treatment  of  both  sexes  are  alike,  seems  to  prove  that  the 
"higher  male  death-rate  is  an  impressed,  natural,  and  constitutional 
"peculiarity  due  to  sex  alone." 

52  'West  Riding  Lunatic  Asylum  Reports,'  vol.  i.  1871,  p.  8.  Sir  J. 
Simpson  has  proved  that  the  head  of  the  male  infant  exceeds  that 
of  the  female  by  3-8ths  of  an  inch  in  circumference,  and  by  l-8th  in 
transverse  diameter.  Quetelet  has  shown  that  woman  is  born  smaller 
than  man;    see  Dr.  Duncan,  'Fecundity,  Fertihty,  Sterility,'  1871,  p.  382. 

53  With  the  savage  Guaranys  of  Paraguay,  according  to  the  accu- 
rate Azara  ('Voyages  dans  I'Amerique  merid.'  tom.  ii.  1809,  p.  60,  179), 
the  women  are  to  the  men  in  the  proportion  of  14  to  13. 


PROPORTION   OF    THE    SEXES.  239 

excess  of  male  over  female  births  is  less  when  they  are  illegitimate 
than  when  legitimate."*  This  has  been  explained  by  different 
writers  in  many  different  ways,  as  from  the  mothers  being  gener- 
ally young,  from  the  large  proportion  of  first  pregnancies,  &c. 
But  we  have  seen  that  male  infants,  from  the  large  size  of  their 
heads,  suffer  more  than  female  infants  during  parturition;  and 
as  the  mothers  of  illegitimate  children  must  be  more  liable 
than  other  women  to  undergo  bad  labors,  from  various  causes, 
such  as  attempts  at  concealment  by  tight  lacing,  hard  work,  dis- 
tress of  mind,  &c.,  their  male  infants  would  proportionally  suffer. 
And  this  probably  is  the  most  efficient  of  all  the  causes  of  the 
proportion  of  males  to  females  born  alive  being  less  amongst 
illegitimate  children  than  amongst  the  legitimate.  With  most 
animals  the  greater  size  of  the  adult  male  than  of  the  female, 
is  due  to  the  stronger  males  having  conquered  the  weaker  in 
their  struggles  for  the  possession  of  the  females,  and  no  doubt 
it  is  owing  to  this  fact  that  the  two  sexes  of  at  least  some  ani- 
mals differ  in  size  at  birth.  Thus  we  have  the  curious  fact  that 
we  may  attribute  the  more  frequent  deaths  of  male  than  female 
infants,  especially  amongst  the  illegitimate,  at  least  in  part  to 
sexual  selection. 

It  has  often  been  supposed  that  the  relative  age  of  the  two 
parents  determines  the  sex  of  the  offspring;  and  Prof.  Leuckart^^ 
has  advanced  what  he  considers  sufficient  evidence,  with  respect 
to  man  and  certain  domesticated  animals,  that  this  is  one  im- 
portant though  not  the  sole  factor  in  the  result.  So  again  the 
period  of  impregnation  relatively  to  the  state  of  the  female  has 
been  thought  by  some  to  be  the  efficient  cause;  but  recent  observa- 
tions discountenance  this  belief.  According  to  Dr.  Stockton- 
Hough,^°  the  season  of  the  year,  the  poverty  or  wealth  of  the 
parents,  residence  in  the  country  or  in  cities,  the  crossing  of 
foreign  immigrants,  &c.,  all  influence  the  proportion  of  the  sexes. 
With  mankind,  polygamy  has  also  been  supposed  to  lead  to  the 
birth  of  a  greater  proportion  of  female  infants;  but  Dr.  J.  Camp- 
belP'  carefully  attended  to  this  subject  in  the  harems  of  Siam, 
and  concludes  that  the  proportion  of  male  to  female  births  is  the 
same  as  from  monogamous  unions.  Hardly  any  animal  has  been 
rendered  so  highly  polygamous  as  the  English  race-horse,  and 
we  shall  immediately  see  that  his  male  and  female  offspring  are 
almost  exactly  equal  in  number.  I  will  now  give  the  facts  which 
I  have  collected  with  respect  to  the  proportional  numbers  of  the 

^  Babbage,  'Edinburgh  Journal  of  Science,'  1829,  vol.  i.  p.  88;  also 
p.  90,  on  still-born  children.  On  illegitimate  children  in  England,  see 
'Report  of   Registrar-General   for  1866,'   p.   xv. 

ss  Leuckart  in  Wagner  'Handworterbuch  der  Phys.'  B.  iv.  1853,  s.  774. 

^  Social  Science  Assoc,   of  Philadelphia,  1874. 

6"^  'Anthropological  Review,'  April,  1870,   p.   cvili. 


240  THE   DESCENT  OF  MAN. 

sexes  of  various  animals;    and  will  then  briefly  discuss  how  far 
selection  has  come  into  play  in  determining  the  result. 

Horses,  — Mr.  Tegetmeier  has  been  so  kind  as  to  tabulate  for  me 
from  the  'Racing  Calendar'  the  births  of  race-horses  during  a 
period  of  twenty-one  years,  viz.,  from  1846  to  1867;  1849  being 
omitted,  as  no  returns  were  that  year  published.  The  total  births 
were  25,560,^^^  consisting  of  12,763  males  and  12,797  females,  or  in 
the  proportion  of  99.7  males  to  100  females.  As  these  numbers 
are  tolerably  large,  and  as  they  are  drawn  from  all  parts  of 
England,  during  several  years,  we  may  with  much  confidence 
conclude  that  with  the  domestic  horse,  or  at  least  with  the  race- 
horse, the  two  sexes  are  produced  in  almost  equal  numbers.  The 
fluctuations  in  the  proportions  during  successive  years  are  closely 
like  those  which  occur  with  mankind,  when  a  small  and  thinly- 
populated  area  is  considered;  thus  in  1856  the  male  horses  were 
as  107.1,  and  in  1867  as  only  92.6  to  100  females.  In  the  tabulated 
returns  the  proportions  vary  in  cycles,  for  the  males  exceeded  the 
females  during  six  successive  years;  and  the  females  exceeded  the 
males  during  two  periods  each  of  four  years:  this,  however,  may 
be  accidental;  at  least  I  can  detect  nothing  of  the  kind  with  man 
in  the  decennial  table  in  the  Registrar's  Report  for  1866. 

Dogs.  — During  a  period  of  twelve  years,  from  1857  to  1868,  the 
births  of  a  large  number  of  greyhounds,  throughout  England, 
were  sent  to  the  'Field'  newspaper;  and  I  am  again  indebted  to 
Mr.  Tegetmeier  for  carefully  tabulating  the  results.  The  recorded 
births  were  6878,  consisting  of  3605  males  and  3273  females,  that 
is,  in  the  proportion  of  110.1  males  to  100  females.  The  greatest 
fluctuations  occurred  in  1864,  when  the  proportion  was  as  95.3 
males,  and  in  1867,  as  116.3  males  to  100  females.  The  above 
average  proportion  of  110.1  to  100  is  probably  nearly  correct  in 
the  case  of  the  greyhound,  but  whether  it  would  hold  with  other 
domesticated  breeds  is  in  some  degree  doubtful.  Mr.  Cupples 
has  inquired  from  several  great  breeders  of  dogs,  and  finds  that 
all  without  exception  believe  that  females  are  produced  in  excess; 
but  he  suggests  that  this  belief  may  have  arisen  from  females 
being  less  valued,  and  from  the  consequent  disappointment  pro- 
ducing a  stronger  impression  on  the  mind. 

^  During  eleven  years  a  record  was  kept  of  the  number  of  mares 
which  proved  barren  or  prematurely  slipped  their  foals;  and  it  de- 
serves notice,  as  showing  how  infertile  these  highly-nurtured  and 
rather  closely-interbred  animals  have  become,  that  not  far  from  one- 
third  of  the  mares  failed  to  produce  living  foals.  Thus  during  1866, 
809  male  colts  and  816  female  colts  were  bom,  and  743  mares  failed  to 
produce  ofEspring.  During  1867,  836  males  and  902  females  were  born, 
and  794  mares  failed. 


PROPORTION   OF   THE    SEXES.  241 

Sheep. — The  sexes  of  sheep  are  not  ascertained  by  agriculturists 
until  several  months  after  birth,  at  the  period  when  the  males 
are  castrated;  so  that  the  following  returns  do  not  give  the  pro- 
portions at  birth.  Moreover,  I  find  that  several  great  breeders 
in  Scotland,  who  annually  raise  some  thousand  sheep,  are  firmly 
convinced  that  a  larger  proportion  of  males  than  of  females  die 
during  the  first  year  or  two.  Therefore  the  proportion  of  males 
would  be  somewhat  larger  at  birth  than  at  the  age  of  castration. 
This  is  a  remarkable  coincidence  with  what,  as  we  have  seen, 
occurs  with  mankind,  and  both  cases  probably  depend  on  the 
same  cause.  I  have  received  returns  from  four  gentlemen  in 
England  who  have  bred  Lowland  sheep,  chiefly  Leicesters,  during 
the  last  ten  to  sixteen  years;  they  amount  altogether  to  8965 
births,  consisting  of  4407  males  and  4558  females;  that  is  in  the 
proportion  of  96.7  males  to  100  females.  With  respect  to  Cheviot 
and  black  faced  sheep  bred  in  Scotland,  I  have  received  returns 
from  six  breeders,  two  of  them  on  a  large  scale,  chiefly  for  the 
years  1867-1869,  but  some  of  the  returns  extend  back  to  1862. 
The  total  number  recorded  amounts  to  50,685,  consisting  of  25,071 
males  and  25,614  females,  or  in  the  proportion  of  97.9  males  to 
100  females.  If  we  take  the  English  and  Scotch  returns  together, 
the  total  number  amounts  to  59,650,  consisting  of  29,478  males 
and  30,172  females,  or  as  97.7  to  100.  So  that  with  sheep  at  the 
age  of  castration  the  females  are  certainly  in  excess  of  the  males, 
but  probably  this  would  not  hold  good  at  birth.^^ 

Of  Cattle  I  have  received  returns  from  nine  gentlemen  of  982 
births,  too  few  to  be  trusted;  these  consisted  of  477  bull-calves 
and  505  cow-calves;  i.  e.,  in  the  proportion  of  94.4  males  to  100 
females.  The  Rev.  "W.  D.  Pox  informs  me  that  in  1867  out  of  34 
calves  born  on  a  farm  in  Derbyshire  only  one  was  a  bull.  Mr. 
Harrison  Weir  has  inquired  from  several  breeders  of  Pigs,  and 
most  of  them  estimate  the  male  to  the  female  births  as  about  7 
to  6.  This  same  gentleman  has  bred  Rabbits  for  many  years, 
and  has  noticed  that  a  far  greater  number  of  bucks  are  produced 
than  does.    But  estimations  are  of  little  value. 

Of  mammalia  in  a  state  of  nature  I  have  been  able  to  learn  very 
little.  In  regard  to  the  common  rat,  I  have  received  conflicting 
statements.  Mr.  R.  Elliot,  of  Laighwood,  informs  me  that  a  rat- 
catcher assured  him  that  he  had  always  found  the  males  in  great 

s»  I  am  much  indebted  to  Mr.  Cupples  for  having-  procured  for  me 
the  above  returns  from  Scotland  as  well  as  some  of  the  following- 
returns  on  cattle.  Mr.  R.  Elliot,  of  Laighwood,  first  called  my  at- 
tention to  the  premature  deaths  of  the  males,— a  statement  subse- 
quently confirmed  by  Mr.  Aitchison  and  others.  To  this  latter  gen- 
tleman, and  to  Mr.  Payan,  I  owe  my  thanks  for  larg-e  returns  as 
to  sheep. 
17 


242  THE  DESCENT  OF  MAN. 

excess,  even  with  the  young  in  the  nest.  In  consequence  of  this, 
Mr.  Elliot  himself  subsequently  examined  some  hundred  old  ones, 
and  found  the  statement  true.  Mr.  F.  Buckland  has  bred  a  large 
number  of  white  rats,  and  he  also  believes  that  the  males  greatly 
exceed  the  females.  In  regard  to  Moles,  it  is  said  that  "the  males 
"are  much  more  numerous  than  the  females;"^''  and  as  the  catch- 
ing of  these  animals  is  a  special  occupation,  the  statement  may 
perhaps  be  trusted.  Sir  A.  Smith,  in  describing  an  antelope  of 
S.  Africa''^  (Kobus  ellipsiprymnus),  remarks,  that  in  the  herds 
of  this  and  other  species,  the  males  are  few  in  number  compared 
with  the  females:  the  natives  believe  that  they  are  born  in  this 
proportion;  others  believe  that  the  younger  males  are  expelled 
from  the  herds,  and  Sir  A.  Smith  says,  that  though  he  has  himself 
never  seen  herds  consisting  of  young  males  alone,  others  af- 
firm that  this  does  occur.  It  appears  probable  that  the  young 
when  expelled  from  the  herd,  would  often  fall  a  prey  to  the  many 
beasts  of  prey  of  the  country. 

BIRDS. 

With  respect  to  the  Fowl,  I  have  received  only  one  account, 
namely,  that  out  of  1001  chickens  of  a  highly-bred  stock  of  Coch- 
ins, reared  during  eight  years  by  Mr.  Stretch,  487  proved  males 
and  514  females;  i.  e.,  as  94.7  to  100.  In  regard  to  domestic 
pigeons  there  is  good  evidence  either  that  the  males  are  produced 
in  excess,  or  that  they  live  longer;  for  these  birds  invariably 
pair,  and  single  males,  as  Mr.  Tegetmeier  informs  me,  can  always 
be  purchased  cheaper  than  females.  Usually  the  two  birds  reared 
from  the  two  eggs  laid  in  the  same  nest  are  a  male  and  a  female; 
but  Mr.  Harrison  Weir,  who  has  been  so  large  a  breeder,  says 
that  he  has  often  bred  two  cocks  from  the  same  nest,  and  seldom 
two  hens;  moreover,  the  hen  is  generally  the  weaker  of  the  two, 
and  more  liable  to  perish. 

With  respect  to  birds  in  a  state  of  nature,  Mr.  Gould  and  oth- 
ers*'^ are  convinced  that  the  males  are  generally  the  more  numer- 
ous; and  as  the  young  males  of  many  species  resemble  the  fe- 
males, the  latter  would  naturally  appear  to  be  the  more  numerous. 
Large  numbers  of  pheasants  are  reared  by  Mr.  Baker  of  Leaden- 
hall  from  eggs  laid  by  wild  birds,  and  he  informs  Mr.  Jenner 
Weir  that  four  or  five  males  to  one  female  are  generally  pro- 
duced.   An  experienced  observer  remarks,^  that  in  Scandinavia 


60  Bell,   'History  of  British  Quadrupeds,'  p.  100, 

61  'Illustrations  of  the  Zoology  of  S.  Africa,'  1849,  pi.  29. 

62  Brehm  ('Illust.  Thierleben,*  B.  iv.  s.  990)  comes  to  the  same  con- 
clusion. 

63  On  the  authority  of  L.    Lloyd,   'Game  Birds  of  Sweden,'  1867,  pp.  12, 
132. 


PROPORTION   OF    THE    SEXES.  243 

the  broods  of  the  capercailzie  and  black-cock  contain  more  males 
than  females;  and  that  with  the  Dal-ripa  (a  kind  of  ptarmigan) 
more  males  than  females  attend  the  leks  or  places  of  courtship; 
but  this  latter  circumstance  is  accounted  for  by  some  observers 
by  a  greater  number  of  hen  birds  being  killed  by  vermin.  From 
various  facts  given  by  White  of  Selborne,"^  it  seems  clear  that  the 
males  of  the  partridge  must  be  in  considerable  excess  in  the  south 
of  England;  and  I  have  been  assured  that  this  is  the  case  in 
Scotland.  Mr.  Weir  on  Inquiring  from  the  dealers,  who  receive 
at  certain  seasons  large  numbers  of  ruffs  (Machetes  pugnax),  was 
told  that  the  males  are  much  the  more  numerous.  This  same 
naturalist  has  also  inquired  for  me  from  the  birdcatchers,  who 
annually  catch  an  astonishing  number  of  various  small  species 
a/live  for  the  London  market,  and  he  was  unhesitatingly  answered 
by  an  old  and  trustworthy  man,  that  with  the  chaffinch  the  males 
are  in  large  excess;  he  thought  as  high  as  2  males  to  1  female, 
or  at  least  as  high  as  5  to  3.*^^  The  males  of  the  blackbird,  he 
likewise  maintained,  were  by  far  the  more  numerous,  whether 
caught  by  traps  or  by  netting  at  night.  These  statements  may 
apparently  be  trusted,  because  this  same  man  said  that  the  sexes 
are  about  equal  with  the  lark,  the  twite  (Linaria  montana),  and 
goldfinch.  On  the  other  hand,  he  is  certain  that  with  the  com- 
mon linnet,  the  females  preponderate  greatly,  but  unequally  during 
different  years;  during  some  years  he  has  found  the  females  to 
the  males  as  four  to  one.  It  shoald,  however,  be  borne  in  mind, 
that  the  chief  season  for  catching  birds  does  not  begin  till  Sep- 
tember, so  that  with  some  species  partial  migrations  may  have 
begun,  and  the  flocks  at  this  period  often  consist  of  hens  alone. 
Mr.  Salvin  paid  particular  attention  to  the  sexes  of  the  humming- 
birds in  Central  America,  and  he  is  convinced  that  with  ihost 
of  the  species  the  males  are  in  excess;  thus  one  year  he  procured 
204  specimens  belonging  to  ten  species,  and  these  consisted  of  166 
males  and  of  only  38  females.  With  two  other  species  the  females 
were  in  excess:  but  the  proportions  apparently  vary  either  dur- 
ing different  seasons  or  in  different  localities;  for  on  one  occasion 
the  males  of  Campylopterus  hemileucurus  were  to  the  females  as 
5  to  2,  and  on  another  occasion^^  in  exactly  the  reversed  ratio.    As 

«4  'Nat.  Hist,  of  Selborne,'  letter  xxix.  edit,  of  1825,  vol.  i.  p.  139. 

^  Mr.  Jenner  Weir  received  similar  information,  on  making  in- 
quiries during  the  following  year.  To  show  the  number  of  living 
chaffinches  caught,  I  may  mention  that  in  1869  there  was  a  match  be- 
tween two  experts,  and  one  man  caught  in  a  day  62,  and  another  40, 
male  ahaffinches.  The  greatest  number  ever  caught  by  one  man  in 
a  single  day  was  70. 

66  'Ibis,'  vol.  ii.  p.  260,  as  quoted  in  Gould's  'Trochilidae,'  1861,  p.  52. 
For  the  foregoing  proportions,  I  am  indebted  to  Mr.  Salvin  for  a 
table  of  his  results. 


244  THE  DESCENT  OF  MAN. 

bearing  on  this  latter  point,  I  may  add,  that  Mr.  Powys  found 
in  Corfu  and  Epirus  the  sexes  of  the  chaffinch  keeping  apart,  and 
"the  females  by  far  the  most  numerous;"  whilst  in  Palestine  Mr. 
Tristram  found  "the  male  flocks  appearing  greatly  to  exceed  the 
"female  in  number.""  So  again  with  the  Quiscalus  major,  Mr.  G.  - 
Taylor'^  says,  that  in  Florida  there  were  "very  few  females  in 
"proportion  to  the  males,"  whilst  in  Honduras  the  proportion  was 
the  other  way,  the  species  there  having  the  character  of  a  polyg- 
amist. 

FISH. 

With  Fish  the  proportional  numbers  of  the  sexes  can  be  ascer- 
tained only  by  catching  them  in  the  adult  or  nearly  adult  state; 
and  there  are  many  difficulties  in  arriving  at  any  just  conclu- 
sion.*'^ Infertile  females  might  readily  be  mistaken  for  males, 
as  Dr.  Giinther  has  remarked  to  me  in  regard  to  trout.  With  some 
species  the  males  are  believed  to  die  soon  after  fertilizing  the 
ova.  With  many  species  the  males  are  of  much  smaller  size  than 
the  females,  so  that  a  large  number  of  males  would  escape  from 
the  same  net  by  which  the  females  were  caught.  M.  Carbonnier,'"' 
who  has  especially  attended  to  the  natural  history  of  the  pike 
(Esox  lucius),  states  that  many  males,  owing  to  their  small  size, 
are  devoured  by  the  larger  females;  and  he  believes  that  the 
males  of  almost  all  fish  are  exposed  from  this  same  cause  to 
greater  danger  than  the  females.  Nevertheless,  in  the  few  cases 
in  which  the  proportional  numbers  have  been  actually  observed, 
the  males  appear  to  be  largely  in  excess.  Thus  Mr.  R.  Buist,  the 
superintendent  of  the  Stormontfield  experiments,  says  that  in 
1865  out  of  70  salmon  first  landed  for  the  purpose  of  obtaining 
the  ova,  upwards  of  60  were  males.  In  1867  he  again  "calls  atten- 
"tion  to  the  vast  disproportion  of  the  males  to  the  females.  We 
"had  at  the  outset  at  least  ten  males  to  one  female."  Afterwards 
females  sufficient  for  obtaining  ova  were  procured.  He  adds, 
"from  the  great  proportion  of  the  males,  they  are  constantly 
"fighting  and  tearing  each  other  on  the  spawning-beds,"^^  This 
disproportion,  no  doubt,  can  be  accounted  for  in  part,  but  whether 
wholly  is  doubtful,  by  the  males  ascending  the  rivers  before  the 
females.    Mr.  F.  Buckland  remarks  in  regard  to  trout,  that  "it  is 


w  'Ibis,'  1860,  p.  137;  and  1867,  p.  369. 

88  'Ibis,'  1862,  p.  137. 

«9L.euckart  quotes  Bloch  (Wagner,  'Handworterbuch  der  Phys.'  B. 
iv.  1853,  s.  775),  that  with  fish  there  are  twice  as  many  males  as 
females. 

''o  Quoted  in  the  'Farmer,'  March  18,  1869,  p.  369. 

"  'The  Stormontfield  Piscicultural  Experiments,'  1866,  p.  23.  The 
•Field'  newspaper,  June  29th,  1867. 


*  PROPORTION    OF    THE    SEXES.  246 

"a  curious  fact  that  the  males  preponderate  very  largely  in  num- 
"ber  over  the  females.  It  invariably  happens  that  when  the  first 
"rush  of  fish  is  made  to  the  net,  there  will  be  at  least  seven  or 
"eight  males  to  one  female  found  captive,  I  cannot  quite  account 
"for  this;  either  the  males  are  more  numerous  than  the  females, 
"or  the  latter  seek  safety  by  concealment  rather  than  flight."  He 
then  adds,  that  by  carefully  searching  the  banks  sufficient  fe- 
males for  obtaining  ova  can  be  found."^  Mr.  H.  Lee  informs  me 
that  out  of  212  trout,  taken  for  this  purpose  in  Lord  Ports- 
mouth's park,  150  were  males  and  62  females. 

The  males  of  the  Cyprinidae  likewise  seem  to  be  in  excess;  but 
several  members  of  this  Family,  viz.,  the  carp,  tench,  bream  and 
minnow,  appear  regularly  to  follow  the  practice,  rare  in  the  ani- 
mal kingdom,  of  polyandry;  for  the  female  while  spawning  is 
always  attended  by  two  males,  one  on  each  side,  and  in  the  case 
of  the  bream  by  three  or  four  males.  This  fact  is  so  well  known, 
that  it  is  always  recommended  to  stock  a  pond  with  two  male 
tenches  to  one  female,  or  at  least  with  three  males  to  two  females. 
With  the  minnow,  an  excellent  observer  states,  that  on  the  spawn- 
ing-beds the  males  are  ten  times  as  numerous  as  the  females; 
when  a  female  comes  amongst  the  males,  "she  is  immediately 
"pressed  closely  by  a  male  on  each  side;  and  when  they  have 
"been  in  that  situation  for  a  time,  are  superseded  by  other  two 
"males.  "^^ 

INSECTS. 

In  this  great  Class,  the  Lepidoptera  almost  alone  afford  means 
for  judging  of  the  proportional  numbers  of  the  sexes;  for  they 
have  been  collected  with  special  care  by  many  good  observers, 
and  have  been  largely  bred  from  the  egg  or  caterpillar  state.  I 
had  hoped  that  some  breeders  of  silk-moths  might  have  kept  an 
exact  record,  but  after  writing  to  France  and  Italy,  and  consult- 
ing various  treatises,  I  cannot  find  that  this  has  ever  been  done. 
The  general  opinion  appears  to  be  that  the  sexes  are  nearly 
equal,  but  in  Italy,  as  I  hear  from  Professor  Canestrini,  many 
breeders  are  convinced  that  the  females  are  produced  in  excess. 
This  same  naturalist,  however,  informs  me,  that  in  the  two  yearly 
broods  of  the  Ailanthus  silk-moth  (Bombyx  cynthia),  the  males 
greatly  preponderate  in  the  first,  whilst  in  the  second  the  two 
sexes  are  nearly  equal,  or  the  females  rather  in  excess. 

In  regard  to  Butterflies  in  a  state  of  nature,  several  observers 

"  'Land  and  Water,'  1868,  p.  41. 

'^Yarrell,   'Hist.  British  Pishes,'  vol.  1.  1826,  p.  307;   on  the  Cyprinus 
carpio,  p.  331;    on  the  Tinea  vulgaris,  p.  331;    on  the  Abramis  brama, 
p.  336.     See,  for  the  minnow  (Leuciscus  phoxinus),   'Loudon's  Mag.  of 
Nat.  Hist.'  vol.  v.  1832,  p.  682. 
17 


246  THE  DESCENT  OF  MAN. 

have  been  much  struck  by  the  apparently  enormous  preponder- 
ance of  the  males.^*    Thus  Mr.  Bates/'^  in  speaking  of  several  spe- 
cies, about  a  hundred  in  number,  which  inhabit  the  Upper  Ama- 
zons, says  that  the  males  are  much  more  numerous  that  the  fe- 
males, even  in  the  proportion  of  a  hundred  to  one.     In  North 
America,  Edwards,  who  had  great  experience,  estimates  in  the 
genus  Papilio  the  males  to  the  females  as  four  to  one;    and  Mr. 
Walsh,  who  informed  me  of  this  statement,  says  that  with  P. 
turnus  this  is  certainly  the  case.    In  South  Africa,  Mr.  R.  Trimen 
found  the  males  in  excess  in  19  species  ;'*'  and  in  one  of  these, 
which  swarms  in  open  places,  he  estimated  the  number  of  males 
as  fifty  to  one  female.    With  another  species,  in  which  the  males 
are  numerous  in  certain  localities,  he  collected  only  five  females 
during  seven  years.    In  the  island  of  Bourbon,  M.  Maillard  states 
that  the  males  of  one  species  of  Papilio  are  twenty  times  as  nu- 
merous as  the  females."    Mr.  Trimen  informs  me  that  as  far  as  he 
has  himself  seen,  or  heard  from  others,  it  is  rare  for  the  females 
of  any  butterfly  to  exceed  the  males  in  number;  but  three  South 
African  species  perhaps  offer  an  exception.    Mr.  Wallace^*  states 
that  the  females  of  Ornithoptera  croesus,  in  the  Malay  archipelago, 
are  more  common  and  more  easily  caught  than  the  males;   but  this 
is  a  rare  butterfly,  I  may  here  add,  that  in  Hyperythra,  a  genus  of 
moths,  Guenee  says,  that  from  four  to  five  females  are  sent  in 
collections  from  India  for  one  male. 

When  this  subject  of  the  proportional  numbers  of  the  sexes  of 
insects  was  brought  before  the  Entomological  Society,^^  it  was 
generally  admitted  that  the  males  of  most  Lepidoptera,  in  the 
adult  or  imago  state,  are  caught  in  greater  numbers  than  the 
females:  but  this  fact  was  attributed  by  various  observers  to  the 
more  retiring  heabits  of  the  females,  and  to  the  males  emerging 
earlier  from  the  cocoon.  This  latter  circumstance  is  well  known 
to  occur  with  most  Lepidoptera,  as  well  as  with  other  insects.  So 
that  as  M.  Personnat  remarks,  the  males  of  the  domesticated 
Bombyx  Yamamai,  are  useless  at  the  beginning  of  the  season,  and 
the  females  at  the  end,  from  the  want  of  mates.^*  I  cannot,  how- 
ever, persuade  myself  that  these  causes  suffice  to  explain  the 


'*Lieuckart  quotes  Meinecke  (Wagner,  'Handworterbuch  der  Phys.' 
B.  iv.  1853,  s.  775)  that  the  males  of  Butterflies  are  three  or  four  times 
as  numerous  as  the  females. 

'5  'The  Naturalist  on  the  Amazons,*  vol.  ii.  1863,  pp.  228,  347. 

'8  Four  of  these  cases  are  given  by  Mr.  Trimen  in  his  'Rhopalocera 
Africae  Australis.' 

"  Quoted  by  Trimen,  'Transact.  Ent.  Soc.,'  vol.  v.  part  iv.  1866,  p.  330. 

'8  'Transact.  Linn.  Soc.,'  vol.  xxv.  p.  37. 

7»  'Proc.  Entomolog.  Soc.,'  Feb.  17th,  1868. 

80  Quoted  by  Dr.  Wallace  in  'Proc.  Ent.  Soc.,'  3rd  series,  vol.  v.  1867. 
p.  487. 


PROPORTION   OF    THE    SEXES.  247 

great  exeess  of  males,  in  the  above  cases  of  certain  butterflies 
which  are  extremely  common  in  their  native  countries.  Mr.  Stain- 
ton,  who  has  paid  very  close  attention  during  many  years  to  the 
smaller  moths,  informs  me  that  when  he  collected  them  in  the 
imago  state,  he  thought  that  the  males  were  ten  times  as  numerous 
as  the  females,  but  that  since  he  has  reared  them  on  a  large  scale 
from  the  caterpillar  state,  he  is  convinced  that  the  females  are 
the  more  numerous.  Several  entomologists  concur  in  this  view. 
Mr.  Doubleday,  however,  and  some  others,  take  an  opposite  view, 
and  are  convinced  that  they  have  reared  from  the  eggs  and  cater- 
pillars a  larger  proportion  of  males  than  of  females. 

Besides  the  more  active  habits  of  the  males,  their  earlier  emerg- 
ence from  the  cocoon,  and  in  some  cases  their  frequenting  more 
open  stations,  other  causes  may  be  assigned  for  an  apparent  or 
real  difference  in  the  proportional  numbers  of  the  sexes  of  Lepi- 
doptera,  when  captured  in  the  imago  state,  and  when  reared  from 
the  egg  or  caterpillar  state.  I  hear  from  Professor  Canestrini, 
that  it  is  believed  by  many  breeders  in  Italy,  that  the  female  cat- 
erpillar of  the  silk-moth  suffers  more  from  the  recent  disease  than 
the  male;  and  Dr.  Staudlinger  informs  me  that  in  rearing  Lepi- 
doptera  more  females  die  in  the  cocoon  than  males.  With  many 
species  the  female  caterpillar  is  larger  than  the  male,  and  a  collect- 
or would  naturally  choose  the  finest  specimens,  and  thus  uninten- 
tionally collect  a  larger  number  of  females.  Three  collectors  have 
told  me  that  this  was  their  practice;  but  Dr.  Wallace  is  sure  that 
most  collectors  take  all  the  specimens  which  they  can  find  of  the 
rarer  kinds,  which  alone  are  worth  the  trouble  of  rearing.  Birds 
when  surrounded  by  caterpillars  would  probably  devour  the  larg- 
est; and  Professor  Canestrini  informs  me  that  in  Italy  some 
breeders  believe,  though  on  insuSacient  evidence,  that  in  the  first 
broods  of  the  Ailanthus  silk-moth,  the  wasps  destroy  a  larger 
number  of  the  female  than  of  the  male  caterpillars.  Dr.  Wal- 
lace further  remarks  that  female  caterpillars,  from  being  larger 
than  the  males,  require  more  time  for  their  development,  and 
consume  more  food  and  moisture;  and  thus  they  would  be  ex- 
posed during  a  longer  time  to  danger  from  ichneumons,  birds, 
&c.,  and  in  times  of  scarcity  would  perish  in  greater  numbers. 
Hence  it  appears  quite  possible  that  in  a  state  of  nature,  fewer 
female  Lepidoptera  may  reach  maturity  than  males;  and  for 
our  special  object  we  are  (concerned  with  their  relative  numbers 
at  maturity,  when  the  sexes  are  ready  to  propagate  their  kind. 

The  manner  in  which  the  males  of  certain  moths  congregate  in 
extraordinary  numbers  round  a  single  female,  apparently  indi- 
cates a  great  excess  of  males,  though  this  fact  may  perhaps  be 
accounted  for  by  the  earlier  emergence  of  the  males  from  their 
cocoons.  Mr.  Stainton  informs  me  that  from  twelve  to  twenty 
males,  may  often  be  seen  congregated  round  a  female  Elachista 


248  THE  DESCENT  OF  MAN. 

rufocinerea.  It  is  well  known  that  if  a  virgin  Lasiocampa  qiiercus 
or  Saturnia  carpini  be  exposed  in  a  cage,  vast  nuiibers  of  males 
collect  round  her,  and  if  confined  in  a  room  will  even  come  down 
the  chimney  to  her.  Mr.  Doubleday  believes  that  he  has  seen 
from  fifty  to  a  hundred  males  of  both  these  species  attracted  in  the 
course  of  a  single  day  by  a  female  in  confinement.  In  the  Isle 
of  Wight  Mr.  Trimen  exposed  a  box  in  which  a  female  of  the 
Lasiocampa  had  been  confined  on  the  previous  day,  and  five  males 
soon  endeavored  to  gain  admittance.  In  Australia,  M.  Verreaux, 
having  placed  the  female  of  a  small  Bombyx  in  a  t30x  in  his  pock- 
et, was  followed  by  a  crowd  of  males,  so  that  about  200  entered 
the  house  with  him.^^ 

Mr.  Doubleday  has  called  my  attention  to  M.  Staudinger's^^  list 
of  Lepidoptera,  which  gives  the  prices  of  the  males  and  females 
of  300  species  or  well-marked  varieties  of  butterflies  (Rhopalo- 
cera).  The  prices  for  both  sexes  of  the  very  common  species  are 
of  course  the  same;  but  in  114  of  the  rarer  species  they  differ; 
the  males  being  in  all  cases,  excepting  one,  the  cheaper.  On  an 
average  of  the  prices  of  the  113  species,  the  price  of  the  male  to 
that  of  the  female  is  as  100  to  149;  and  this  apparently  indicates 
that  inversely  the  males  exceed  the  females  in  the  same  propor- 
tion. About  2000  species  or  varieties  of  moths  (Heterocera)  are 
catalogued,  those  with  wingless  females  being  here  excluded  on 
account  of  the  difference  in  habits  between  the  two  sexes:  of 
these  2000  species,  141  differ  in  price  according  to  sex,  the  males 
of  130  being  cheaper,  and  those  of  only  11  being  dearer  than  the 
females.  The  average  price  of  the  males  of  the  130  species,  to 
that  of  the  females,  is  as  100  to  143.  With  respect  to  the  butter- 
flies in  this  priced  list,  Mr.  Doubleday  thinks  (and  no  man  in 
England  has  had  more  experience),  that  there  is  nothing  in  the 
habits  of  these  species  which  can  account  for  the  difference  in 
the  prices  of  the  two  sexes,  and  that  it  can  be  accounted  for  only 
by  an  excess  in  the  number  of  the  males.  But  I  am  bound  to 
add  that  Dr.  Staudinger  informs  me,  that  he  is  himself  of  a  dif- 
ferent opinion.  He  thinks  that  the  less  active  habits  of  the  fe- 
males and  the  earlier  emergence  of  the  males  will  account  for 
his  collectors  securing^  a  larger  number  of  males  than  of  females, 
and  consequently  for  the  lower  prices  of  the  former.  With  respect 
to  specimens  reared  from  the  caterpillar-state.  Dr.  Staudinger 
believes,  as  previously  stated,  that  a  greater  number  of  females 
than  of  males  die  whilst  confined  in  the  cocoons.  He  adds  that 
with  certain  species  one  sex  seems  to  preponderate  over  the  other 
during  certain  years. 

Of  direct  observations    on    the    sexes   of  Lepidoptera  reared 

^  Blanchard,   'Metamorphoses,  Moeurs  des  Insectes/  1868,  pp.  225-226. 
•»  'Lepidopteren-Doubletten    Liste,'    Berlin,    No.    x.    1866. 


PROPORTION   OF    THE    SEXES.  249 

either  from  eggs  or  caterpillars,  I  have  received  only  the  few 
following  cases:  — 

Males.    Females. 
The  Rev.  J.  Hellins^  of  Exeter  reared,  during  1868, 

images  of  73  species,  which  consisted  of 153  137 

Mr.  Albert  Jones  of  Eltham  reared,  during  1868, 

imagos  of  9  species,  which  consisted  of 159  126 

During  1869  he  reared  imagos  from  4  species,  con- 
sisting  of    114  112 

Mr.  Buckler    of    Emsworth,    Hants,  during   1869, 

reared  imagos  from  74  species,  consisting  of 180  169 

Dr.  Wallace  of  Colchester  reared  from  one  brood 

of  Bombyx  cynthia 52  48 

Dr.  Wallace  raised,  from  cocoons  of  Bombyx  Pern- 

yi  sent  from  China,  during  1869 224  123 

Dr.   Wallace   raised,   during  1868   and   1869,   from 

two  lots  of  cocoons  of  Bombyx  yama-mai 52  46 

Total 934  761 

So  that  in  these  eight  lots  of  cocoons  and  eggs,  males  were  pro- 
duced in  excess.  Taken  together  the  proportion  of  males  is  as 
122.7  to  100  females.  But  the  numbers  are  hardly  large  enough 
to  be  trustworthy. 

On  the  whole,  from  these  various  sources  of  evidence,  all  point- 
ing in  the  same  direction,  I  infer  that  with  most  species  of  Lepi- 
doptera,  the  mature  males  generally  exceed  the  females  in  num- 
ber, whatever  the  proportions  may  be  at  their  first  emergence 
from  the  egg. 

With  reference  to  the  other  Orders  of  insects,  I  have  been  able 
to  collect  very  little  reliable  information.  With  the  stag-beetle 
(Lucanus  cervus)  "the  males  appear  to  be  much  more  numerous 
"than  the  females;"  but  when,  as  Cornelius  remarked  during  1867, 
an  unufeual  number  of  these  beetles  appeared  in  one  part  of  Ger- 
many, the  females  appeared  to  exceed  the  males  as  six  to  one. 
With  one  of  the  Elateridse,  the  males  are  said  to  be  much  more 
numerous  than  the  females,  and  "two  or  three  are  often  found 
"united  with  one  female  ;^^  so  that  here  polyandry  seems  to  pre- 
"vail."    With  Siagonium  (StaphylinidsB),  in  which  the  males  are 

83  This  naturalist  has  been  so  kind  as  to  send  me  some  results  from 
former  years,  in  which  the  females  seemed  to  preponderate;  but  so 
many  of  the  figures  were  estimates,  that  I  found  it  impossible  to 
tabulate  them. 

^  Gunther's  'Record  of  Zoological  Literature,'  1867,  p.  260.  On  the 
excess  of  female  Lucanus,  ibid.  p.  250.  On  the  males  of  Lucanus  in 
England,  Westwood,  'Modern  Class,  of  Insects,'  vol.  i.  p.  187.  On  the 
Siagonium,  ibid.  p.  172. 


250  THE  DESCENT  OF  MAN. 

furnished  with  horDS,  "the  females  are  far  more  numerous  than 
"the  opposite  sex."  Mr.  Janson  stated  at  the  Entomological  So- 
ciety that  the  females  of  the  bark-feeding  Tomicus  villosus  are  so 
common  as  to  be  a  plague,  whilst  the  males  are  so  rare  as  to  be 
hardly  known. 

It  is  hardly  worth  while  saying  anything  about  the  proportion 
of  the  sexes  in  certain  species  and  even  groups  of  insects,  for 
the  males  are  unknown  or  very  rare,  and  the  females  are  parthen- 
ogenetic,  that  is,  fertile  without  sexual  union;  examples  of  this 
are  afforded  by  several  of  the  Cynipidae.^^  In  all  the  gall-making 
CynipidaB  known  to  Mr.  Walsh,  the  females  are  four  or  five  times 
as  numerous  as  the  males;  and  so  it  is,  as  he  informs  me,  with 
the  gall-making  Cecidomyiiae  (Diptera).  With  some  common 
species  of  Saw-flies  (Tenthredinae)  Mr.  F.  Smith  has  reared  hun- 
dreds of  specimens  from  larvae  of  all  sizes,  but  has  never,  reared 
a  single  male:  on  the  other  hand,  Curtis  says,^^  that  with  certg-in 
species  (Athalia),  bred  by  him,  the  males  were  to  the  females  as 
six  to  one;  whilst  exactly  the  reverse  occurred  with  the  mature 
insects  of  the  same  species  caught  in  the  fields.  In  the  family  of 
Bees,  Hermann  Miiller"  collected  a  large  number  of  specimens  of 
many  species,  and  reared  others  from  the  cocoons,  and  counted 
the  sexes.  He  found  that  the  males  of  some  species  greatly  ex- 
ceeded the  females  in  number;  in  others  the  reverse  occurred;  and 
in  others  the  two  sexes  were  nearly  equal.  But  as  in  most  cases  the 
males  emerge  from  the  cocoons  before  the  females,  they  are  at 
the  commencement  of  the  breeding  season  practically  in  excess. 
Miiller  also  observed  that  the  relative  number  of  the  two  sexes 
in  some  species  differed  much  in  different  localities.  But  as  H. 
Miiller  has  himself  remarked  to  me,  these  remarks  must  be  re- 
ceived with  some  caution,  as  one  sex  might  more  easily  escape  ob- 
servation than  the  other.  Thus  his  brother  Fritz  Miiller  has 
noticed  in  Brazil  that  the  two  sexes  of  the  same  species  of  bee 
sometimes  frequent  different  kinds  of  flowers.  With  respect  to 
the  Orthoptera,  I  know  hardly  anything  about  the  relative  num- 
ber of  the  sexes:  Korte,^^  however,  says  that  out  of  500  locusts 
which  he  examined,  the  males  were  to  the  females  as  five  to  six. 
With  the  Neuroptera,  Mr.  Walsh  states  that  in  many,  but  by 
no  means  in  all  the  species  of  the  Odonatous  group,  there  is  a 
great  overplus  of  males:  in  the  genus  Hetserina,  also,  the  males 
are  generally  at  least  four  times  as  numerous  as  the  females. 
In  certain  species  in  the  genus  Gomphus  the  males  are  equally  in 

85  Walsh,   in    'The  American   Entomologist,*    vol.    i.    1869,    p.    103.    F. 
Smith,  'Record  of  Zoological  Literature,'  1867,  p.  328. 
88  'Farm  Insects,'  pp.  45-46. 

87  Anwendung-  der  Darwinschen  Lehre  Verh.  d.  n.  V.  Jahrg.  xxiv.' 

88  'Die  Strich,   Zug  oder  Wanderheuschrecke,'   1828,  p.  20. 


PROPOKTION   OF   THE    SEXES.  251 

excess,  whilst  in  two  other  species,  the  females  are  twice  or 
thrice  as  numerous  as  the  males.  In  some  European  species  of 
Psocus  thousands  of  females  may  he  collected  without  a  single 
male,  whilst  with  other  species  of  the  same  genus  both  sexes  are 
common.^''  In  England,  Mr.  MacLachlan  has  captured  hundreds 
of  the  female  Apatania  muliebris,  but  has  never  seen  the  male; 
and  of  Boreus  hyemalis  only  four  or  five  males  have  been  seen 
here.^°  With  most  of  these  species  (excepting  the  Tenthredinae) 
there  is  at  present  no  evidence  that  the  females  are  subject  to 
parthenogenesis;  and  thus  we  see  how  ignorant  we  are  of  the 
causes  of  the  apparent  discrepancy  in  the  proportion  of  the  two 
sexes. 

In  the  other  Classes  of  the  Articulata  I  have  been  able  to  collect 
still  less  information.  With  Spiders,  Mr.  Blackwall,  who  has 
carefully  attended  to  this  class  during  many  years,  writes  to  me 
that  the  males  from  their  more  erratic  habits  are  more  commonly 
seen,  and  therefore  appear  more  numerous.  This  is  actually  the 
case  with  a  few  species;  but  he  mentions  several  species  in  six 
genera,  in  which  the  females  appear  to  be  much  more  numerous 
than  the  males.^^  The  small  size  of  the  males  in  comparison  with 
the  females  (a  peculiarity  which  is  sometimes  carried  to  an  ex- 
treme degree),  and  their  widely  different  appearance,  may  ac- 
count in  some  instances  for  their  rarity  in  collections.®^ 

Some  of  the  lower  Crustaceans  are  able  to  propagate  their 
kind  asexually,  and  this  will  account  for  the  extreme  rarity  of  the 
males:  thus  Von  Siebold®*  carefully  examined  no  less  than  13,000 
specimens  of  Apus  from  twenty-one  localities,  and  amongst  these 
he  found  only  319  males.  With  some  other  forms  (as  Tanais 
and  Cypris),  as  Fritz  Miiller  informs  me,  there  is  reason  to  be- 
lieve that  the  males  are  much  shorter-lived  than  the  females; 
and  this  would  explain  their  scarcity,  supposing  the  two  sexes  to 
be  at  first  equal  in  number.  On  the  other  hand,  Miiller  has  in- 
variably taken  far  more  males  than  females  of  the  Diastylidae 
and  of  Cypridina  on  the  shores  of  Brazil;  thus  with  a  species  in 
the  latter  genus,  63  specimens  caught  the  same  day  included  57 
males;  but  he  suggests  that  this  preponderance  may  be  due  to 
some  unknown  difference  in  the  habits  of  the  two  sexes.  With 
one  of  the  higher  Brazilian  crabs,  namely  a  Gelasimus,  Fritz 

*»  'Observations  on  N.  American  Neuroptera,'  by  H.  Hagen  and  B. 
D.  Walsh,   'Proc.  Ent.  Soc.  Philadelphia,'  Oct.  1863,  pp.  168,  223,  239. 

^  'Proc.   Ent.   Soc.   London,   Feb.  17,   1868. 

81  An  other  great  authority  with  respect  to  this  class,  Prof.  Thorell 
of  TJpsala  ('On  European  Spiders,'  1869-70,  part  t  p.  205)  speaks  as  If 
female  spiders  were  generally  commoner  than  the  males. 

*2  See,  on  this  subject,  Mr.  O.  P,  Cambridge,  as  quoted  in  'Quarterly 
Journal  of  Science,'  1868,  p.  429. 

93  'Beitrage  zur  Parthenogenesis,'  p.  174. 


252  THE  DESCENT  OF  MAN. 

Miiller  found  the  males  to  be  more  numerous  than  the  females. 
According  to  the  large  experience  of  Mr.  C.  Spence  Bate,  the 
reverse  seems  to  be  the  case  with  six  common  British  crabs,  the 
names  of  which  he  has  given  me. 


The  proportion  of  the  sexes  in  relation  to  natv/ral  selection. 

There  is  reason  to  suspect  that  in  some  cases  man  has  by  selec- 
tion indirectly  influenced  his  own  sex-producing  powers.  Cer- 
tain women  tend  to  produce  during  their  whole  lives  more  chil- 
dren of  one  sex  than  of  the  other:  and  the  same  holds  good  of 
many  animals,  for  instance,  cows  and  horses;  thus  Mr.  Wright 
of  Yeldersley  House  informs  me  that  one  of  his  Arab  mares, 
though  put  seven  times  to  different  horses,  produced  seven  fil- 
lies. Though  I  have  very  little  evidence  on  this  head,  analogy 
would  lead  to  the  belief,  that  the  tendency  to  produce  either 
sex  would  be  inherited  like  almost  every  other  peculiarity,  for 
instance,  that  of  producing  twins;  and  concerning  the  above 
tendency  a  good  authority,  Mr.  J.  Downing,  has  communicated 
to  me  facts  which  seem  to  prove  that  this  does  occur  in  certain 
families  of  short-horn  cattle.  Col.  Marshal?*  has  recently  found 
on  careful  examination  that  the  Todas,  a  hill-tribe  of  India,  con- 
sist of  112  males  and  84  females  of  all  ages — that  is  in  a  ratio  of 
133.3  males  to  100  females.  The  Todas,  who  are  polyandrous  in 
their  marriages,  during  former  times  invariably  practiced  female 
infanticide;  but  this  practice  has  now  been  discontinued  for  a 
considerable  period.  Of  the  children  born  within  late  years,  the 
males  are  more  numerous  than  the  females,  in  the  proportion  of 
124  to  100.  Colonel  Marshall  accounts  for  this  fact  in  the  fol- 
lowing ingenious  manner:  "Let  us  for  the  purpose  of  illustra- 
"tion  take  three  families  as  representing  an  average  of  the 
"entire  tribe;  say  that  one  mother  gives  birth  to  six  daughters 
"and  no  sons;  a  second  mother  has  six  sons  only,  whilst  the 
"third  mother  has  three  sons  and  three  daughters.  The  first 
"mother,  following  the  tribal  custom,  destroys  four  daughters 
"and  preserves  two.  The  second  retains  her  six  sons.  The  third 
"kills  two  daughters  and  keeps  one,  as  also  her  three  sons.  We 
"have  then  from  the  three  families,  nine  sons  and  three  daugh- 
"ters,  with  which  to  continue  the  breed.  But  whilst  the  males 
"belong  to  families  in  which  the  tendency  to  produce  sons  is 
"great,  the  females  are  of  those  of  a  converse  inclination.  Thus 
"the  bias  strengthens  with  each  generation,  until,  as  we  find, 
"families  grow  to  have  habitually  more  sons  than  daughters." 

That  this  result  would  follow  from  the  above  form  of  infanti- 
cide seems  almost  certain;    that  is  if  we  assume  that  a  sex- 

94  «xhe  Todas,'  1873,  pp.  100,  111,  194,  196. 


PROPORTION  OF  THE  SEXES.  253 

producing  tendency  is  inherited.  But  as  the  above  numbers  are 
so  extremely  scanty,  I  have  searched  for  additional  evidence, 
but  cannot  decide  whether  what  I  have  found  is  trustworthy; 
nevertheless  the  facts  are,  perhaps,  worth  giving.  The  Maories 
of  New  Zealand  have  long  practiced  infanticide;  and  Mr.  Fen- 
ton®^  states  that  he  "has  met  with  instances  of  women  who  have 
"destroyed  four,  six,  and  even  seven  children,  mostly  females. 
"However,  the  universal  testimony  of  those  best  qualified  to 
"judge,  is  conclusive  that  this  custom  has  for  many  years  been 
"almost  extinct.  Probably  the  year  1835  may  be  named  as  the 
"period  of  its  ceasing  to  exist."  Now  amongst  the  New  Zea- 
landers,  as  with  the  Todas,  male  births  are  considerably  in  ex- 
cess. Mr.  Fenton  remarks  (p.  30),  "One  fact  is  certain,  although 
"the  exact  period  of  the  commencement  of  this  singular  condi- 
"tion  of  the  disproportion  of  the  sexes  cannot  be  demonstratively 
"fixed,  it  is  quite  clear  that  this  course  of  decrease  was  in  full 
"operation  during  the  years  1830  to  1844,  when  the  non-adult 
"population  of  1844  was  being  produced,  and  has  continued  with 
"great  energy  up  to  the  present  time."  The  following  statements 
are  taken  from  Mr.  Fenton  (p.  26),  but  as  the  numbers  are  not 
large,  and  as  the  census  was  not  accurate,  uniform  results  cannot 
be  expected.  It  should  be  borne  in  mind  in  this  and  the  follow- 
ing cases,  that  the  normal  state  of  every  population  is  an  excess 
of  women,  at  least  in  all  civilized  countries,  chiefly  owing  to  the 
greater  mortality  of  the  male  sex  during  youth,  and  partly  to  acci- 
dents of  all  kinds  later  in  life.  In  1858,  the  native  population  of 
New  Zealand  was  estimated  as  consisting  of  31,667  males  and 
24,303  females  of  all  ages,  that  is  in  the  ratio  of  130.3  males  to 
100  females.  But  during  this  same  year,  and  in  certain  limited 
districts,  the  numbers  were  ascertained  with  much  care,  and  the 
males  of  all  ages  were  here  753  and  the  females  616;  that  is  in 
the  ratio  of  122.2  males  to  100  females.  It  is  more  important 
for  us  that  during  this  same  year  of  1858,  the  non-adult  males 
within  the  same  district  were  found  to  be  178,  and  the  non-adult 
females  142,  that  is  in  the  ratio  of  125.3  to  100.  It  may  be  added 
that  in  1844,  at  which  period  female  infanticide  had  only  lately 
ceased,  the  non-adult  males  in  one  district  were  281,  and  the 
non-adult  females  only  194,  that  is  in  the  ratio  of  144.8  males  to 
100  females. 

In  the  Sandwich  Islands,  the  males  exceed  the  females  in 
number.  Infanticide  was  formerly  practiced  there  to  a  frightful 
extent,  but  was  by  no  means  confined  to  female  infants,  as  is 
shown  by  Mr.  Ellis,^^  and  as  I  have  been  informed  by  Bishop 

05  'Aboriginal    Inhabitants    of    New    Zealand;    Government    Report,' 
1859,  p.  36. 
88  'Narrative  of  a  Tour  through  Hawaii,'  1826,  p.  298. 


254  THE  DESCENT  OF  MAN. 

Staley  and  the  Rev.  Mr.  Coan.  Nevertheless,  another  apparently 
trustworthy  writer,  Mr.  Jarves,*^  whose  observations  apply  to 
the  whole  archipelago,  remarks: — "Numbers  of  women  are  to 
"be  found,  who  confess  to  the  murder  of  from  three  to  six  or 
"eight  children;"  and  he  adds,  "females  from  being  considered 
"less  useful  than  males  were  more  often  destroyed."  From  what 
is  known  to  occur  in  other  parts  of  the  world,  this  statement  is 
probable;  but  must  be  received  with  much  caution.  The  prac- 
tice of  infanticide  ceased  about  the  year  1819,  when  idolatry  was 
abolished  and  missionaries  settled  in  the  Islands.  A  careful  cen- 
sus in  1839  of  the  adult  and  taxable  men  and  women  in  the  island 
of  Kauai  and  in  one  district  of  Oahu  (Jarves,  p.  404),  gives  4723 
males  and  3776  females;  that  is  in  the  ratio  of  125.08  to  100.  At 
the  same  time  the  number  of  males  under  fourteen  years  in  Kauai 
and  under  eighteen  in  Oahu  was  1797,  and  of  females  of  the  same 
ages  1429;  and  here  we  have  the  ratio  of  125.75  males  to  100 
females. 

In  a  census  of  all  the  islands  in  1850,®^  the  males  of  all  ages 
amount  to  86,272,  and  the  females  to  33,128,  or  as  109.49  to  100. 
The  males  under  seventeen  years  amounted  to  10,773,  and  the 
females  under  the  same  age  to  9593,  or  as  112.3  to  100.  From  the 
census  of  1872,  the  proportion  of  males  of  all  ages  (including 
half-castes)  to  females,  is  as  125.36  to  100.  It  must  be  borne  in 
mind  that  all  these  returns  for  the  Sandwich  Islands  give  the 
proportion  of  living  males  to  living  females,  and  not  of  the  births; 
and  judging  from  all  civilized  countries  the  proportion  of  males 
would  have  been  considerably  higher  if  the  numbers  had  referred 
to  births.^ 

;    87  'History  of  the  Sandwich  Islands,'  1843,  p,  93. 

»8  This  is  given  in  the  Rev.  H.  T.  Cheever's  'Life  in  tlie  Sandwich 
Islands,'  1851,  p.  277. 

»» Dr.  Coulter,  in  describing  ('Journal  R.  Geograph.  Soe,'  vol.  v. 
1835,  p.  67)  the  state  of  California  about  the  year  1830,  says  that  the 
natives,  reclaimed  by  the  Spanish  missionaries,  have  nearly  all  per- 
ished, or  are  perishing,  although  well  treated,  not  driven  from  their 
native  land,  and  kept  from  the  use  of  spirits.  He  attributes  this,  in 
great  part,  to  the  undoubted  fact  that  the  men  greatly  exceed  the 
wom.en  in  number;  but  he  does  not  know  whether  this  is  due  to  a 
failure  of  female  offspring,  or  to  more  females  dying  during  early 
youth.  The  latter  alternative,  according  to  all  analogy,  is  very  im- 
probable. He  adds  that  "infanticide,  properly  so  called,  is  not  com- 
"mon,  though  very  frequent  recourse  is  had  to  abortion."  If  Dr. 
Coulter  is  correct  about  infanticide,  this  case  cannot  be  advanced  in 
support  of  Col.  Marshall's  view.  From  the  rapid  decrease  of  the  re- 
claimed natives,  we  may  suspect  that,  as  in  the  cases  lately  given, 
their  fertility  has  been  diminished  from  changed  habits  of  life. 

I  had  hoped  to  gain  some  light  on  this  subject  from  the  breeding  of 
do&s;  inasmuch  as  in  most  breeds,  with  the  exception,  perhaps,  of 


PROPOKTION  OF  THE  SEXES.  255 

From  the  several  foregoing  cases  we  have  some  reason  to 
believe  that  infanticide  practiced  in  the  manner  above  explained, 
tends  to  make  a  male-producing  race;  but  I  am  far  from  sup- 
posing that  this  practice  in  the  case  of  man,  or  some  analogous 
process  with  other  species,  has  been  the  sole  determining  cause 
of  an  excess  of  males.  There  may  be  some  unknown  law  leading 
to  this  result  in  decreasing  races,  which  have  already  become 
somewhat  infertile.  Besides  the  several  causes  previously  al- 
luded to,  the  greater  facility  of  parturition  amongst  savages, 
and  the  less  consequent  injury  to  their  male  infants,  would 
tend  to  increase  the  proportion  of  live-born  males  to  females. 
There  does  not,  however,  seem  to  be  any  necessary  connection 
between  savage  life  and  a  marked  excess  of  males;  that  is  if  we 
may  judge  by  the  character  of  the  scanty  offspring  of  the  lately 
existing  Tasmanians  and  of  the  crossed  offspring  of  the  Tahi- 
tians  now  inhabiting  Norfolk  Island. 

As  the  males  and  females  of  many  animals  differ  somewhat  in 
habits  and  are  exposed  in  different  degrees  to  danger,  it  is 
probable  that  in  many  cases,  more  of  one  sex  than  of  the  other 
are  habitually  destroyed.  But  as  far  as  I  can  trace  out  the  com- 
plication of  causes,  an  indiscriminate  though  large  destruction 
of  either  sex  would  not  tend  to  modify  the  sex-producing  power 
of  the  species.  With  strictly  social  animals,  such  as  bees  or  ants, 
which  produce  a  vast  number  of  sterile  and  fertile  females  in 
comparison  with  the  males,  and  to  whom  this  preponderance  is 
of  paramount  importance,  we  can  see  that  those  communities 
would  flourish  best  which  contained  females  having  a  strong 
inherited  tendency  to  produce  more  and  more  females;  and  in 
such  cases  an  unequal  sex-producing  tendency  would  be  ulti- 
mately gained  through  natural  selection.  With  animals  living 
in  herds  or  troops,  in  which  the  males  come  to  the  front  and 
defend  the  herd,  as  with  the  bisons  of  North  America  and  certain 

greyhounds,  many  more  female  puppies  are  destroyed  than  males, 
just  as  with  the  Toda  infants.  Mr.  Cupples  assures  me  that  this  is 
usual  with  Scotch  deer-hounds.  Unfortunately,  I  know  nothing  of 
the  proportion  of  the  sexes  in  any  breed,  excepting  greyhounds,  and 
there  the  male  births  are  to  the  female  as  110.1  to  100.  Now  from  in- 
quiries made  from  many  breeders,  it  seems  that  the  females  are  in 
some  respects  more  esteemed,  though  otherwise  troublesome;  and  it 
does  not  appear  that  the  female  puppies  of  the  best-bred  dogs  are 
systematically  destroyed  more  than  the  males,  though  this  does  some- 
times take  place  to  a  limited  extent.  Therefore  I  am  unable  to  de- 
cide whether  we  can,  on  the  above  principles,  account  for  the  pre- 
ponderance of  male  births  in  greyhounds.  On  the  other  hand,  we 
have  seen  that  with  horses,  cattle,  and  sheep,  which  are  too  valuable 
for  the  young  of  either  sex  to  be  destroyed,  if  there  is  any  difference, 
the  females  are  slightly  in  excess. 


256  THE  DESCENT  OF  MAN. 

baboons,  it  is  conceivable  that  a  male-producing  tendency  might 
be  gained  by  natural  selection;  for  the  individuals  of  the  better 
defended  herds  would  leave  more  numerous  descendants.  In 
the  case  of  mankind  the  advantage  arising  from  having  a  pre- 
ponderance of  men  in  the  tribe  is  supposed  to  be  one  chief  cause 
of  the  practice  of  female  infanticide. 

In  no  case,  as  far  as  we  can  see,  would  an  inherited  tendency 
to  produce  both  sexes  in  equal  numbers  or  to  produce  one  sex 
in  excess,  be  a  direct  advantage  or  disadvantage  to  certain  in- 
dividuals more  than  to  others;  for  instance,  an  individual  with 
a  tendency  to  produce  more  males  than  females  would  not  succeed 
better  in  the  battle  for  life  than  an  individual  with  an  opposite 
tendency;  and  therefore  a  tendency  of  this  kind  could  not  be 
gained  through  natural  selection.  Nevertheless,  there  are  certain 
animals  (for  instance,  fishes  and  cirripedes)  in  which  two  or  more 
males  appear  to  be  necessary  for  the  fertilization  of  the  female; 
and  the  males  accordingly  largely  preponderate,  but  it  is  by  no 
means  obvious  that  this  male-producing  tendency  could  have  been 
acquired.  I  formerly  thought  that  when  a  tendency  to  produce 
the  two  sexes  in  equal  numbers  was  advantageous  to  the  species, 
it  would  follow  from  natural  selection,  but  I  now  see  that  the 
whole  problem  is  so  intricate  that  it  is  safer  to  leave  its  solution 
for  the  future. 


SEXUAL  SELECTION.  257 


CHAPTER  IX. 

SECONDARY    SEXUAL    CHARACTERS    IN   THE    LOWER 
CLASSES    OF    THE    ANIMAL    KINGDOM. 

These  characters  absent  in  the  lower  classes— Brilliant  colors— Mol- 
lusca— Annelids — Crustacea,  secondary  sexual  characters  strongly 
developed;  dimorphism;  color;  characters  not  acquired  before 
maturity— Spiders,  sexual  colors  of;  stridulation  by  the  males— 
Myrlapoda. 

With  animals  belonging  to  the  lower  classes,  the  two  sexes 
are  not  rarely  united  in  the  same  individual,  and  therefore  sec- 
ondary sexual  characters  cannot  be  developed.  In  many  cases 
where  the  sexes  are  separate,  both  are  permanently  attached 
to  some  support,  and  the  one  cannot  search  or  struggle  for  the 
other.  Moreover  it  is  almost  certain  that  these  animals  have  too 
imperfect  senses  and  much  too  low  mental  powers,  to  appreciate 
each  other's  beauty  or  other  attractions,  or  to  feel  rivalry. 

Hence  in  these  clases  or  sub-kingdoms,  such  as  the  Protozoa, 
Coelenterata,  Echinodermata,  Scolecida,  secondary  sexual  charac- 
ters, of  the  kind  which  we  have  to  consider,  do  not  occur;  and 
this  fact  agrees  with  the  belief  that  such  characters  in  the  higher 
classes  have  been  acquired  through  sexual  selection,  which  de- 
pends on  the  will,  desire,  and  choice  of  either  sex.  Nevertheless 
some  few  apparent  exceptions  occur;  thus,  as  I  hear  from  Dr. 
Baird,  the  males  of  certain  Entozoa,  or  internal  parasitic  worms, 
differ  slightly  in  color  from  the  females;  but  we  have  no  reason 
to  suppose  that  such  differences  have  been  augmented  through 
sexual  selection.  Contrivances  by  which  the  male  holds  the  fe- 
male, and  which  are  indispensable  for  the  propagation  of  the 
species,  are  independent  of  sexual  selection,  and  have  been  ac- 
quired through  ordinary  selection. 

Many  of  the  lower  animals,  whether  hermaphrodites  or  with 
separate  sexes,  are  ornamented  with  the  most  brilliant  tints,  or 
are  shaded  and  striped  in  an  elegant  manner;  for  instance,  many 
corals  and  sea-anemones  (Actinise),  some  jelly-fish  (Medusae, 
Porpita,  &c.),  some  Planariae,  many  star-fishes,  Echini,  Ascidians, 
&c.;  but  we  may  conclude  from  the  reasons  already  indicated, 
18 


258  THE  DESCENT  OF  MAN. 

namely  the  union  of  the  two  sexes  in  some  of  these  animals,  the 
permanently  affixed  condition  of  others,  and  the  low  mental 
powers  of  all,  that  such  colors  do  not  serve  as  a  sexual  attraction, 
and  have  not  been  acquired  through  sexual  selection.  It  should 
be  borne  in  mind  that  in  no  case  have  we  sufficient  evidence  that 
colors  have  been  thus  acquired,  except  where  one  sex  is  much 
more  brilliantly  or  conspicuously  colored  than  the  other,  and 
where  there  is  no  difference  in  habits  between  the  sexes  sufficient 
to  account  for  their  different  colors.  But  the  evidence  is  rendered 
as  complete  as  it  can  ever  be,  only  when  the  more  ornamented 
individuals,  almost  always  the  males,  voluntarily  display  their 
attractions  before  the  other  sex;  for  we  cannot  believe  that  such 
display  is  useless,  and  if  it  be  advantageous,  sexual  selection 
will  almost  inevitably  follow.  We  may,  however,  extend  this 
conclusion  to  both  sexes,  when  colored  alike,  if  their  colors  are 
plainly  analogous  to  those  of  one  sex  alone  in  certain  other 
species  of  the  same  group. 

How,  then,  are  we  to  account  for  the  beautiful  or  even  gor- 
geous colors  of  many  animals  in  the  lowest  classes?  It  appears 
doubtful  whether  such  colors  often  serve  as  a  protection;  but 
that  we  may  easily  err  on  this  head,  will  be  admitted  by  every 
one  who  reads  Mr.  Wallace's  excellent  essay  on  this  subject.  It 
would  not,  for  instance,  at  first  occur  to  any  one  that  the  trans- 
parency of  the  Medusae,  or  jelly-fishes,  is  of  the  highest  service 
to  them  as  a  protection;  but  when  we  are  reminded  by  Hackel 
that  not  only  the  medusse,  but  many  floating  mollusca,  crusta- 
ceans, and  even  small  oceanic  fishes  partake  of  this  same  glass- 
like appearance,  often  accompanied  by  prismatic  colors,  we  can 
hardly  doubt  that  they  thus  escape  the  notice  of  pelagic  birds 
and  other  enemies.  M.  Giard  is  also  convinced^  that  the  bright 
tints  of  certain  sponges  and  ascidians  serve  as  a  protection.  Con- 
spicuous colors  are  likewise  beneficial  to  many  animals  as  a 
warning  to  their  would-be  devourers  that  they  are  distasteful,  or 
that  they  possess  some  special  means  of  defense;  but  this  subject 
will  be  discussed  more  conveniently  hereafter. 

We  can,  in  our  ignorance  of  most  of  the  lowest  animals,  only 
say  that  their  bright  tints  result  either  from  the  chemical  na- 
ture or  the  minute  structure  of  their  tissues,  independently  of  any 
benefit  thus  derived.  Hardly  any  color  is  finer  than  that  of  ar- 
terial blood;  but  there  is  no  reason  to  suppose  that  the  color  of 
the  blood  is  in  itself  any  advantage;  and  though  it  adds  to  the 
beauty  of  the  maiden's  cheek,  no  one  will  pretend  that  it  has  been 
acquired  for  this  purpose.  So  again  with  many  animals,  espe- 
cially the  lower  ones,  the  bile  is  richly  colored;  thus,  as  I  am  in- 
formed by  Mr.  Hancock,  the  extreme  beauty  of  the  Eolidse  (naked 

1  'Archives  de  Zoolog-.    Exper.,'   Oct.   1872,  p.  563. 


MOLLUSCS.  259 

sea-slugs)  is  chiefly  due  to  the  biliary  glands  being  seen  through 
the  translucent  integuments — this  beauty  being  probably  of  no 
service  to  these  animals.  The  tints  of  the  decaying  leaves  in  an 
American  forest  are  described  by  every  one  as  gorgeous;  yet  no 
one  supposes  that  these  tints  are  of  the  least  advantage  to  the 
trees.  Bearing  in  mind  how  many  substances  closely  analogous 
to  natural  organic  compounds  have  been  recently  formed  by  chem- 
ists, and  which  exhibit  the  most  splendid  colors,  it  would  have 
been  a  strange  fact  if  substances  similarly  colored  had  not  often 
originated,  independently  of  any  useful  end  thus  gained,  in  the 
complex  laboratory  of  living  organisms. 

The  sub-kingdom  of  the  Mollusca.  — Throughout  this  great  divis- 
ion of  the  animal  kingdom,  as  far  as  I  can  discover,  secondary 
sexual  characters,  such  as  we  are  here  considering,  never  occur. 
Nor  could  they  be  expected  in  the  three  lowest  classes,  namely  in 
the  Ascidians,  Polyzoa,  and  Brachiopods  (constituting  the  Mollus- 
coida  of  some  authors),  for  most  of  these  animals  are  permanently 
affixed  to  a  support  or  have  their  sexes  united  in  the  same  Individ' 
ual.  In  the  Lamellibranchiata,  or  bivalve  shells,  hermaphrodit- 
ism is  not  rare.  In  the  next  higher  class  of  the  Gasteropoda,  or 
univalve  shells,  the  sexes  are  either  united  or  separate.  But  in 
the  latter  case  the  males  never  possess  special  organs  for  find- 
ing, securing,  or  charming  the  females,  or  for  fighting  with  other 
males.  As  I  am  informed  by  Mr.  Gwyn  Jeffreys,  the  sole  external 
difference  between  the  sexes  consists  in  the  shell  sometimes  dif- 
fering a  little  in  form;  for  instance,  the  shell  of  the  male  peri- 
winkle (Littorina  littorea)  is  narrower  and  has  a  more  elongated 
spire  than  that  of  the  female.  But  differences  of  this  nature,  it 
may  be  presumed,  are  directly  connected  with  the  act  of  reproduc- 
tion, or  with  the  development  of  the  ova. 

The  Gasteropoda,  though  capable  of  locomotion  and  furnished 
with  imperfect  eyes,  do  not  appear  to  be  endowed  with  sufficient 
mental  powers  for  the  members  of  the  same  sex  to  struggle  to- 
gether in  rivalry,  and  thus  to  acquire  secondary  sexual  characters. 
Nevertheless  with  the  pulmoniferous  gasteropods,  or  land-snails, 
the  pairing  is  preceded  by  courtship;  for  these  animals,  though 
hermaphrodites,  are  compelled  by  their  structure  to  pair  together. 
Agassiz  remarks,-  "Quiconque  a  eu  I'occasion  d'observer  les  amours 
"des  limagons,  ne  saurait  mettre  en  doute  le  seduction  deployee 
"dans  les  mouvements  et  les  allures  qui  pr6parent  et  accomplis- 
"sent  le  double  embrassement  de  ces  hermaphrodites."  These 
animals  appear  also  susceptible  of  some  degree  of  permanent 
attachment:  an  accurate  obser/er,  Mr.  Lonsdale,  informs  me 
that  he  placed  a  pair  of  land-snails  (Helix  pomatia),  one  of  which 

2  'D©  I'Espece  et  de  la  Class.  &c.,  1869,  p.  106. 


260  THE   DESCENT  OF  MAN. 

was  weakly,  into  a  small  and  ill-provided  garden.  After  a  short 
time  the  strong  and  healthy  individual  disappeared,  and  was 
traced  by  its  track  of  slime  over  a  wall  into  an  adjoining  well- 
stocked  garden.  Mr.  Lonsdale  concluded  that  it  had  deserted  its 
sickly  mate;  but  after  an  absence  of  twenty-four  hours  it  re- 
turned, and  apparently  communicated  the  result  of  its  successful 
exploration,  for  both  then  started  along  the  same  track  and  dis- 
appeared over  the  wall. 

Even  in  the  highest  class  of  the  Mollusca,  the  Cephalopoda  or 
cuttlefishes,  in  which  the  sexes  are  separate,  secondary  sexual 
characters  of  the  present  kind  do  not,  as  far  as  I  can  discover, 
occur.  This  is  a  surprising  circumstance,  as  these  animals  pos- 
sess highly-developed  sense-organs  and  have  considerable  mental 
powers,  as  will  be  admitted  by  every  one  who  has  watched  their 
artful  endeavors  to  escape  from  an  enemy.^  Certain  Cephalopoda, 
however,  are  characterized  by  one  extraordinary  sexual  character, 
namely,  that  the  male  element  collects  within  one  of  the  arms 
or  tentacles,  which  is  then  cast  off,  and  clinging  by  its  sucking- 
discs  to  the  female,  lives  for  a  time  an  independent  life.  So  com- 
pletely does  the  cast-off  arm  resemble  a  separate  animal,  that  it 
was  described  by  Cuvier  as  a  parasitic  worm  under  the  name  of 
Hectocotyle.  But  this  marvelous  structure  may  be  classed  as  a 
primary  rather  than  as  a  secondary  sexual  character. 

Although  with  the  Mollusca  sexual  selection  does  not  seem  to 
have  come  into  play;  yet  many  univalve  and  bivalve  shells,  such 
as  volutes,  cones,  scallops,  &c.,  are  beautifully  colored  and  shaped. 
The  colors  do  not  appear  in  most  cases  to  be  of  any  use  as  a 
protection;  they  are  probably  the  direct  result,  as  in  the  lowest 
classes,  of  the  nature  of  the  tissues;  the  patterns  and  the  sculp- 
ture of  the  shell  depending  on  its  manner  of  growth.  The  amount 
of  light  seems  to  be  influential  to  a  certain  extent;  for  although, 
as  repeatedly  stated  by  Mr.  Gwyn  Jeffreys,  the  shells  of  some  spe- 
cies living  at  a  profound  depth  are  brightly  colored,  yet  we  gen- 
erally see  the  lower  surfaces,  as  well  as  the  parts  covered  by 
the  mantle,  less  highly-colo-'ed  than  the  upper  and  exposed  sur- 
faces.* In  some  cases,  as  with  shells  living  amongst  corals  or 
brightly-tinted  sea- weeds,  the  bright  colors  may  serve  as  a  pro- 
tection.^   But  that  many  of  the  nudi-branch  mollusca,  or  sea.-slugs, 

3  See,  for  instance,  the  account  which  I  have  given  in  my  'Journal 
of  Researches,'  1845,  p.   7. 

*  I  have  given  ('Geolog.  Observations  on  Volcanic  Islands,'  1844,  p.  53) 
a  curious  instance  of  the  influence  of  light  on  the  colors  of  a  frondes- 
cent  incrustation,  deposited  by  the  surf  on  the  coast-rocks  of  Ascen- 
sion, and  formed  by  the  solution  of  triturated  sea-shells. 

^  Dr.  Morse  has  lately  discussed  this  subject  in  his  paper  on  the 
Adaptive  Coloration  of  Mollusca,  'Proc.  Boston  See.  of  Nat.  Hist.,' 
vol.  xiv.,  April,  1871. 


MOLLUSCS.  261 

are  as  beautifully  colored  as  any  shells,  may  be  seen  in  Messrs. 
Alder  and  Hancock's  magnificent  work;  and  'from  information 
kindly  given  me  by  Mr.  Hancock,  it  seems  extremely  doubtful 
whether  these  colors  usually  serve  as  a  protection.  With  some 
species  this  may  be  the  case,  as  with  one  kind  which  lives  on  the 
green  leaves  of  algae,  and  is  itself  bright-green.  But  many  bright- 
ly-colored, white  or  otherwise  conspicuous  species,  do  not  seek 
concealment;  whilst  again  some  equally  conspicuous  species,  as 
well  as  other  dull-colored  kinds,  live  under  stones  and  in  dark 
recesses.  So  that  with  these  nudi-branch  molluscs,  color  appar- 
ently does  not  stand  in  any  close  relation  to  the  nature  of  the 
places  which  they  inhabit. 

These  naked  sea-slugs  are  hermaphrodites,  yet  they  pair  to- 
gether, as  do  land-snails,  many  of  which  have  extremely  pretty 
shells.  It  is  conceivable  that  two  hermaphrodites,  attracted  by 
each  other's  greater  beauty,  might  unite  and  leave  offspring  which 
would  inherit  their  parents'  greater  beauty.  But  with  such  lowly- 
organized  creatures  this  is  extremely  improbable.  Nor  is  it  at  all 
obvious  how  the  offspring  from  the  more  beautiful  pairs  of  her- 
maphrodites would  have  any  advantage  over  the  offspring  of  the 
less  beautiful,  so  as  to  increase  in  number,  unle-ss  indeed  vigor 
and  beauty  generally  coincided.  V/e  have  not  here  the  case  of  a 
number  of  males  becoming  mature  before  the  females,  with  the 
more  beautiful  males  selected  by  the  more  vigorous  females.  If, 
indeed,  brilliant  colors  were  beneficial  to  a  hermaphrodite  ani- 
mal in  relation  to  its  general  habits  of  life,  the  more  brightly- 
tinted  individuals  would  succeed  best  and  would  increase  in 
number;  but  this  would  be  a  case  of  natural  and  not  of  sexual 
selection. 

Sub-Mngdom  of  the  Vermes:  Class,  Annelida  (or  Sea-worms). 
— In  this  class,  although  the  sexes,  when  separate,  sometimes  dif- 
fer from  each  other  in  characters  of  such  importance  that  they 
have  been  placed  under  distinct  genera  or  even  families,  yet  the 
differences  do  not  seem  of  the  kind  which  can  be  safely  at- 
tributed to  sexual  selection.  These  animals  are  often  beautifully 
colored,  but  as  the  sexes  do  not  differ  in  this  respect,  we  are  but 
little  concerned  with  them.  Even  the  Nemertians,  though  so 
lowly  organized,  "vie  in  beauty  and  variety  of  coloring  with  any 
"other  group  in  the  invertebrate  series;"  yet  Dr.  Mcintosh^  can- 
not discover  that  these  colors  are  of  any  service.  The  sedentary 
annelids  become  duller-colored,  according  to  M.  Quatrefages,' 
after  the  period  of  reproduction;  and  this  I  presume  may  be  at- 

«  See  his  beautiful  monograph  on  'British  Annelids,  part  i.  1873,  p.  3. 
'  See    M.    Perrier,    'I'Origine   de   I'Homme   d'apres   Darwin,'    'Revue 
Scientifique,'   Feb.   1873,   p.    866. 
13 


262  THE  DESCENT  OF  MAN. 

tributed  to  tlieir  less  vigorous  condition  at  that  time.  All  these 
worm-like  animals  apparently  stand  too  low  in  the  scale  for  the 
individuals  of  either  sex  to  exert  any  choice  in  selecting  a  part- 
ner, or  for  the  individuals  of  the  same  sex  to  struggle  together 
in  rivalry. 

Sitb-Mngdom  of  the  Arthropoda:  Class,  Crustacea. — In  this 
great  class  we  first  meet  with  undoubted  secondary  sexual  char- 
acters, often  developed  in  a  remarkable  manner.  Unfortunately 
the  habits  of  crustaceans  are  very  imperfectly  known,  and  we 
cannot  explain  the  uses  of  many  structures  peculiar  to  one  sex. 
With  the  lower  parasitic  species  the  males  are  of  small  size,  and 
they  alone  are  furnished  with  perfect  swimming-legs,  antennae 
and  sense-organs;  the  females  being  destitute  of  these  organs, 
with  their  bodies  often  consisting  of  a  mere  distorted  mass.  But 
these  extraordinary  differences  between  the  two  sexes  are  no 
doubt  related  to  their  widely  different  habits  of  life,  and  con- 
sequently do  not  concern  us.  In  various  crustaceans,  belonging 
to  distinct  families,  the  anterior  antennae  are  furnished  with  pe- 
culiar thread-like  bodies,  which  are  believed  to  act  as  smelling- 
organs,  and  these  are  much  more  numerous  in  the  males  than  in 
the  females.  As  the  males,  without  any  unusual  development  of 
their  olfactory  organs,  would  almost  certainly  be  able  sooner  or 
later  to  find  the  females,  the  increased  number  of  the  smelling- 
threads  has  probably  been  acquired  through  sexual  selection,  by 
the  better  provided  males  having  been  the  more  successful  in 
finding  partners  and  in  producing  offspring.  Fritz  Miiller  has 
described  a  remarkable  dimorphic  species  of  Tanais,  in  which  the 
male  is  represented  by  two  distinct  forms,  which  never  graduate 
into  each  other.  In  the  one  form  the  male  is  furnished  with 
more  numerous  smelling-threads,  and  in  the  other  form  with 
more  powerful  and  more  elongated  chelae  or  pincers,  which  serve 
to  hold  the  female.  Fritz  Miiller  suggests  that  these  differences 
between  the  two  male  forms  of  the  same  species  may  have  origi- 
nated in  certain  individuals  having  varied  in  the  number  of  the 
smelling-threads,  whilst  other  individuals  varied  in  the  shape 
and  size  of  their  chelae;  so  that  of  the  former,  those  which  were 
best  able  to  find  the  female,  and  of  the  latter,  those  which  were 
best  able  to  hold  her,  have  left  the  greatest  number  of  progeny 
to  inherit  their  respective  advantages.^ 

In  some  of  the  lower  crustaceans,  the  right  anterior  antenna 
of  the  male  differs  greatly  in  structure  from  the  left,  the  latter 

8  'Facts  and  Arguments  for  Darwin,'  English  translat.  1869,  p.  20. 
See  the  previous  discussion  on  the  olfactory  threads.  Sars  has  de- 
scribed a  somewhat  analogous  case  (as  quoted  in  'Nature,'  1870,  p.  455) 
in  a  Norwegian  crustacean,   the  Pontoporeia  affinis. 


CRUSTACEANS. 


263 


resembling  in  its  simple  tapering  joints  the  antennae  of  the  fe- 
male. In  the  male  the  modified  antenna  is  either  swollen  in  the 
^  middle  or  angularly   bent,  or  converted 

(fig.  4.)  into  an  elegant,  and  sometimes 
wonderfully  complex,  prehensile  organ.' 
It  serves,  as  I  hear  from  Sir  J.  Lubbock, 
to  hold  the  female,  and  for  this  same  pur- 
pose one  of  the  two  posterior  legs  (b)  on 
the  same  side  of  the  body  is  converted 
into  a  forceps.  In  another  family  the 
inferior  or  posterior  antennae  are  "curi- 
ously zigzagged'*  in  the  males  alone. 

In  the  higher  crustaceans  the  anterior 
legs  are  developed  into  chelae  or  pincers; 
and  these  are  generally  larger  in  the  male 
than  in  the  female, — so  much  so  that  the 
market  value  of  the  male  edible  crab  (Can- 
cer pagurus),  according  to  Mr.  C.  Spence 
Bate,  is  five  times  as  great  as  that  of  the 
female.  In  many  species  the  chelae  are  of 
unequal  size  on  the  opposite  side  of  the 
body,  the  right-hand  one  being,  as  I  am 
informed  by  Mr.  Bate,  generally,  though 
not  invariably,  the  largest.  This  inequal- 
ity is  also  often  much  greater  in  the  male 
than  in  the  female.  The  two  chelae  of  the 
male  often  differ  in  structure  (figs.  5,  6, 
and  7), the  smaller  one  resembling  that  of 
the  female.  V/hat  advantage  is  gained  by 
their  inequality  in  size  on  the  opposite 
sides  of  the  body,  and  by  the  inequality 
being  much  greater  in  the  male  than  in 
the  female;  and  why,  when  they  are  of 
equal  size,  both  are  often  much  larger  in  the  male  than  in  the 
female,  is  not  known.  As  I  hear  from  Mr.  Bate,  the 
chelae  are  sometimes  of  such  length  and  size  that  they 
cannot  possibly  be  used  for  carrying  food  to  the  mouth. 
In  the  males  of  certain  fresh-water  prawns  (Palaemon)  the  right 
leg  is  actually  longer  than  the  whole  body.^°  The  great  size  of 
the  one  leg  with  its  chel®  may  aid  the  male  in  fighting  with  his 


Fig-.  4.    Labidocera  Dar- 
winii  (from  Lubbock). 

a.  Part  of  right  anterior 
antenna  of  male,  form- 
ing a  prehensile  organ. 

b.  Posterior  pair  of  thor- 
acic leg's  of  male. 

c.  Ditto  of  female. 


» See  Sir  J.  Lubbock  in  'Annals  and  Mag.  of  Nat.  Hist.,'  vol.  xi. 
1853,  pi.  i.  and  x. ;  and  vol.  xii.  (1853)  pi,  vii.  See,  also,  Lubbock  in 
'Transact.  Ent.  Soc.,'  vol.  iv.  new  series,  1856-1858,  p.  8.  With  respect 
to  the  zig-zagged  antennae  mentioned  below,  see  Fritz  Muller,  'Facts 
and  Arguments  for  Darwin,'  1869,  p.  40,  foot-note. 

">  See  a  paper  by  Mr.  C.  Spence  Bate,  with  figures,  in  'Proc.  Zoolog. 


264 


THE  DESCENT  OF  MAN. 


rivals;   but  this  will  not  account  for  their  inequality  in  the  fe- 
male on  the  opposite  sides  of  the  body.    In  Gelasimus,  according 


Fl*'.  5.    Anterior  part  of  body  of  Callianassa  (from  Milne-Edwards), 

showing  the   unequal  and   differently   constructed   right 

and  left-hand  chelae  of  the  male. 

N.  B.— The  artist  by  mistake  has  reversed  the  drawing,  and  made 
the  left-hand  chela  the  largest. 


Fig.  6. 


Fig.  7. 


Fig.  6.  Second  leg  of  male  OrchestiaTucuratinga  (from  Fritz  Muller^ 
Fig.  7.    Ditto  of  female. 

to  a  statement  quoted  by  Milne-Edwards,"  the  male  and  the  fe- 
male live  in  the  same  burrow,  and  this  shows  that  they  pair; 
the  male  closes  the  mouth  of  the  burrow  with  one  of  its  chel^, 
which  is  enormously  developed;  so  that  here  it  indirectly  serves 
as  a  means  of  defense.    Their  main  use  however,  is  probably  to 


Soc.,'  1868,  p.  363;    and  on  the  nomenclature  of  the  genus,  ibid.  p.  585. 
I  am  greatly  indebted  to  Mr.  Spence  Bate  for  nearly  all  the  above 
statements  with  respect  to  the  chelae  of  the  higher  crustaceans. 
"  'Hist.   Nat.  des  Crust.'  torn.  ii.  1837,  p.  50. 


CRUSTACEANS.  265 

seize  and  to  secure  the  female,  and  this  in  some  instances,  as 
with  G-ammarus,  is  known  to  be  the  case.  The  male  of  the  her- 
mit or  soldier  crab  (Pagurus)  for  weeks  together,  carries  about 
the  shell  inhabited  by  the  female.^-  The  sexes,  however,  of  the 
common  shore-crab  (Carcinus  manas),  as  Mr.  Bate  informs  me, 
unite  directly  after  the  female  has  moulted  her  hard  shell,  when 
she  is  so  soft  that  she  would  be  injured  if  seized  by  the  strong 
pincers  of  the  male;  but  as  she  is  caught  and  carried  about  by 
the  male  before  moulting,  she  could  then  be  seized  with  impunity. 

Fritz  Miiller  states  that  certain  species  of  Melita  are  distin- 
guished from  all  other  amphipods  by  the  females  having  "the 
"coxal  lamellae  of  the  penultimate  pair  of  feet  produced  into 
"hook-like  processes,  of  which  the  males  lay  hold  with  the  hands 
"of  the  first  pair."  The  development  of  these  hook-like  proc- 
esses has  probably  followed  from  those  females  which  were  the 
most  securely  held  during  the  act  of  reproduction,  having  left 
the  largest  number  of  offspring.  Another  Brazilian  amphipod 
(Orchestia  Darwinii,  fig.  8)  presents  a  case  of  dimorphism,  like 
that  of  Tanais;  for  there  are  two  male  forms,  which  differ  in  the 
structure  of  their  chelae.^^  As  either  chela  would  certainly  suf- 
fice to  hold  the  female, — for  both  are  now  used  for  this  purpose, 
— the  two  male  forms  probably  originated  by  some  having  va- 
ried in  one  manner  and  some  in  another;  both  forms  having 
derived  certain  special,  but  nearly  equal  advalitages,  from  their 
differently  shaped  organs. 

It  is  not  known  that  male  crustaceans  fight  together  for  the 
possession  of  the  females,  but  it  is  probably  the  case,  for  with 
most  animals  when  the  male  is  larger  than  the  female,  he  seems 
to  owe  his  greater  size  to  his  ancestors  having  fought  with  other 
males  during  many  generations.  In  most  of  the  orders,  espe- 
cially in  the  highest  or  the  Brachyura,  the  male  is  larger  than  the 
female;  the  parasitic  genera,  however,  in  which  the  sexes  follov/ 
different  habits  of  life,  and  most  of  the  Entomostraca  must  be 
excepted.  The  chelae  of  many  crustaceans  are  weapons  well 
adapted  for  fighting.  Thus  when  a  Devil-crab  (Portunus  puber) 
was  seen  by  a  son  of  Mr.  Bate  fighting  with  a  Carcinus  maenas, 
the  latter  was  soon  thrown  on  its  back,  and  had  every  limb  torn 
from  its  body.  When  several  males  of  a  Brazilian  Gelasimus,  a 
species  furnished  with  immense  pincers,  were  placed  together  in 
a  glass  vessel  by  Fritz  Miiller,  they  mutilated  and  killed  one 
another.  Mr.  Bate  put  a  large  male  Carcinus  masnas  into  a  pan 
of  water,  inhabited  by  a  female  which  was  paired  with  a  smaller 
male;  but  the  latter  was  soon  dispossessed.    Mr.  Bate  adds,  "if 


"  Mr.  C.  Spence  Bate,  'Brit.  Assoc,  Fourth  Report  on  the  Fauna  of 
B.  Devon.' 
13  Fritz  Muller,   'Facts  and  Arguments  for  Darwin,'  1869,  pp.  25-28. 


266 


THE  DESCENT  OF  MAN. 


*'lfiey  fought,  the  victory  was  a  bloodless  one,  for  I  saw  no 
"wounds."  This  same  naturalist  separated  a  male  sand-skipper 
(so  common  on  our  sea-shores),  Gammarus  marinus,  from  its 
female,  both  of  whom  were  imprisoned  in  the  same  vessel  with 
many  individuals  of  the  same  species.    The  female,  when  thus 


Fig.  8.    Orchestia  Darwinii   (from   Fritz  Muller),   showing  the  differ- 
ently-constructed  chelae  of   the   two   male  forms. 

divorced,  soon  joined  the  others.  After  a  time  the  male  was  put 
again  into  the  same  vessel;  and  he  then,  after  swimming  about 
for  a  time,  dashed  into  the  crowd,  and  without  any  fighting  at 
once  took  away  his  wife.  This  fact  shows  that  in  the  Amphi- 
poda,  an  order  low  in  the  scale,  the  males  and  females  recognize 
each  other,  and  are  mutually  attached. 


CRUSTACEANS.  267 

The  mental  powers  of  the  Crustacea  are  probably  higher  than 
at  first  sight  appears  probable.  Any  one  who  tries  to  catch  one 
of  the  shore-crabs,  so  common  on  tropical  coasts,  will  perceive 
how  wary  and  alert  they  are.  There  is  a  large  crab  (Birgus 
latro),  found  on  coral  islands,  which  makes  a  thick  bed  of  the 
picked  fibres  of  the  cocoa-nut,  at  the  bottom  of  a  deep  burrow. 
It  feeds  on  the  fallen  fruit  of  this  tree  by  tearing  off  the  husk, 
fibre  by  fibre;  and  it  always  begins  at  that  end  where  the  three 
eye-like  depressions  are  situated.  It  then  breaks  through  one  of 
these  eyes  by  hammering  with  its  heavy  front  pincers,  and  turn- 
ing round,  extracts  the  albuminous  core  with  its  narrow  posterior 
pincers.  But  these  actions  are  probably  instinctive,  so  that 
they  would  be  performed  as  well  by  a  young  animal  as  by  an  old 
one.  The  following  case,  however,  can  hardly  be  so  considered: 
a  trustworthy  naturalist,  Mr.  Gardner,^*  whilst  watching  a  shore- 
crab  (Gelasimus)  making  its  burrow,  threw  some  shells  towards 
the  hole.  One  rolled  in,  and  three  other  shells  remained  within 
a  few  inches  of  the  mouth.  In  about  five  minutes  the  crab 
brought  out  the  shell  which  had  fallen  in,  and  carried  it  away  to 
the  distance  of  a  foot;  it  then  saw  the  three  other  shells  lying 
near,  and  evidently  thinking  that  they  might  likewise  roll  in, 
carried  them  to  the  spot  where  it  had  laid  the  first.  It  would,  I 
think,  be  difficult  to  distinguish  this  act  from  one  performed  by 
man  by  the  aid  of  reason. 

Mr.  Bate  does  not  know  of  any  well-marked  case  of  difference 
of  color  in  the  two  sexes  of  our  British  crustaceans,  in  which 
respect  the  sexes  of  the  higher  animials  so  often  differ.  In  some 
cases,  however,  the  males  and  females  differ  slightly  in  tint,  but 
Mr.  Bate  thinks  not  more  than  may  be  accounted  for  by  their 
different  habits  of  life,  such  as  by  the  male  wandering  more 
about,  and  being  thus  more  exposed  to  the  light.  Dr.  Power 
tried  to  distinguish  by  color  the  sexes  of  the  several  species  which 
inhabit  the  Mauritius,  but  failed,  except  with  one  species  of 
Squilla,  probably  S.  stylifera,  the  male  of  which  is  described  as 
being  "of  a  beautiful  bluish-green,"  v/ith  some  of  the  appen- 
dages cherry-red,  whilst  the  female  is  clouded  v/ith  brown  and 
gray,  "with  the  red  about  her  much  less  vivid  than  in  the  male."^^ 
In  this  case,  we  may  suspect  the  agency  of  sexual  selection. 
From  M.  Bert's  observations  on  Daphnia,  when  placed  in  a  vessel 
illuminated  by  a  prism,  we  have  reason  to  believe  that  even  the 
lowest  crustaceans  can  distinguish  colors.  With  Saphirina  (an 
oceanic  genus  of  Bntomostraca),  the  males  are  furnished  with 

1*  'Travels  in  the  Interior  of  Brazil,'  1846,  p.  HI.  I  have  given,  in 
my  'Journal  of  Researches,'  p.  463,  an  account  of  the  habits  of  the 
Birgus. 

15  Mr.  Ch.  Fraser,  in  'Proc  Zoolog.  Soc.,'  1869,  p.  3.  I  am  indebted 
to  Mr.  Bate  for  Dr.  Power's  statement. 


268  THE  DESCENT  OF  MAN. 

miuute  shields  or  cell-like  bodies,  which  exhibit  beautiful  chang- 
ing colors;  these  are  absent  in  the  females,  and  in  both  sexes  of 
one  species.^®  It  would,  liowever,  be  extremely  rash  to  conclude 
that  these  curious  organs  serve  to  attract  the  females.  I  am  in- 
formed by  Fritz  Miiller,  that  in  the  female  of  a  Brazilian  species 
of  Gelasimus,  the  whole  body  is  of  a  nearly  uniform  grayish- 
brown.  In  the  male  the  posterior  part  of  the  cephalo-thorax 
is  pure  white,  with  the  anterior  part  of  a  rich  green,  shading 
into  dark  brown;  and  it  is  remarkable  that  these  colors  are  liable 
to  change  in  the  course  of  a  few  minutes — the  white  becoming 
dirty  gray  or  even  black,  the  green  "losing  much  of  its  brilliancy." 
It  deserves  especial  notice  that  the  males  do  not  acquire  their 
bright  colors  until  they  become  mature.  They  appear  to  be 
much  more  numerous  than 'the  females;  they  differ  also  in  the 
larger  size  of  their  chelae.  In  some  species  of  the  genus,  prob- 
ably in  all,  the  sexes  pair  and  inhabit  the  same  burrow.  They 
are  also,  as  we  have  seen,  highly  intelligent  animals.  From 
these  various  considerations  it  seems  probable  that  the  male  in 
this  species  has  become  gaily  ornamented  in  order  to  attract  or 
excite  the  female. 

It  has  just  been  stated  that  the  male  Gelasimus  does  not  ac- 
quire his  conspicuous  colors  until  mature  and  nearly  ready  to 
breed.  This  seems  a  general  rule  in  the  whole  class  in  respect  to 
the  many  remarkable  structural  differences  between  the  sexes. 
We  shall  hereafter  find  the  same  law  prevailing  throughout  the 
great  sub-kingdom  of  the  Vertebrata;  and  in  all  cases  it  is  emi- 
nently distinctive  of  characters  which  have  been  acquired  through 
sexual  selection.  Fritz  Miiller"  gives  some  striking  instances  of 
this  law;  thus  the  male  sand-hopper  (Orchestia)  does  not,  until 
nearly  full  grown,  acquire  his  large  claspers,  which  are  very  dif- 
ferently constructed  from  those  of  the  female;  whilst  young,  his 
claspers  resemble  those  of  the  female. 

Class,  Arachnida  (Spiders). — The  sexes  do  not  generally  differ 
much  in  color,  but  the  males  are  often  darker  than  the  females, 
as  may  be  seen  in  Mr.  Blackwall's  magnificent  work.^^  In  some 
species,  however,  the  difference  is  conspicuous:  thus  the  female 
of  Sparassus  smaragdulus  is  dullish  green,  whilst  the  adult  male 
has  the  abdomen  of  a  fine  yellow,  with  three  longitudinal  stripes 
of  rich  red.  In  certain  species  of  Thomisus  the  sexes  closely  re- 
semble each  other,  in  others  they  differ  much;  and  analogous 
cases  occur  in  many  other  genera.  It  is  often  difficult  to  say 
which  of  the  two  sexes  departs  most  from  the  ordinary  coloration 

^«  Claus,   'Die  freilebenden  Copepoden,'  1863,   s.  35. 
"  'Facts  and  Arguments,'  &c.,  p.  79. 

18  'A  History  of  the  Spiders  of  Great  Britain,'  1861-64.  For  the  fol- 
lowing facts,  see  pp.  77,  88,  102. 


SPIDERS.  269 

of  the  genus  to  whicli  the  species  belong;  but  Mr.  Blackwall 
thinks  that,  as  a  general  rule,  it  is  the  male;  and  Canestrini^^  re- 
marks that  in  certain  genera  the  males  can  be  specifically  dis- 
tinguished with  ease,  but  the  females  with  great  difficulty.  I  am 
informed  by  Mr.  Blackwall  that  the  sexes  whilst  young  usually 
resemble  each  other;  and  both  often  undergo  great  changes  in 
color  during  their  successive  moults,  before  arriving  at  maturity. 
In  other  cases  the  male  alone  appears  to  change  color.  Thus 
the  male  of  the  above  bright-colored  Sparassus  at  first  resembles 
the  female,  and  acquires  his  peculiar  tints  only  when  nearly 
adult.  Spiders  are  possessed  of  acute  senses,  and  exhibit  much 
intelligence;  as  is  well  known,  the  females  often  show  the  strong- 
est affection  for  their  eggs,  which  they  carry  about  enveloped 
in  a  silken  web.  The  males  search  eagerly  for  the  females,  and 
have  been  seen  by  Canestrini  and  others  to  fight  for  possession  of 
them.  This  same  author  says  that  the  union  of  the  two  sexes 
has  been  observed  in  about  twenty  species;  and  he  asserts  posi- 
tively that  the  female  rejects  some  of  the  males  who  court  her, 
threatens  them  with  open  mandibles,  and  at  last  after  long  hesi- 
tation accepts  the  chosen  one.  From  these  several  considera- 
tions, we  may  admit  with  some  confidence  that  the  well-marked 
differences  in  color  between  the  sexes  of  certain  species  are  the 
results  of  sexual  selection;  though  we  have  not  here  the  best 
kind  of  evidence, — the  display  by  the  male  of  his  ornaments. 
From  the  extreme  variability  of  color  in  the  male  of  some  species, 
for  instance  of  Theridion  lineatum,  it  would  appear  that  these 
sexual  characters  of  the  males  have  not  as  yet  become  well  fixed. 
Canestrini  draws  the  same  conclusion  from  the  fact  that  the 
males  of  certain  species  present  two  forms,  differing  from  each 
other  in  the  size  and  length  of  their  jaws;  and  this  reminds  us 
of  the  above  cases  of  dimorphic  crustaceans. 

The  male  is  generally  much  smaller  than  the  female,  some- 
times to  an  extraordinary  degree,==''  and  he  is  forced  to  be  ex- 
tremely cautious  in  making  his  advances,  as  the  female  often 
carries  her  coyness  to  a  dangerous  pitch.  De  Geer  saw  a  male 
that  "in  the  midst  of  his  preparatory  caresses  was  seized  by  the 
"object  of  his  attentions,  enveloped  by  her  in  a  web  and  then  6.e- 

i«  This  author  has  recently  published  a  valuable  essay  on  the  'Car- 
atteri  sessuali  secondarii  degli  Arachnidi,'  in  the  'Atti  della  Soc. 
Veneto-Trentina  di  So.  Nat.  Padova,  vol.  i.  Fasc.  3,  1873. 

20  Aug-.  Vinson  ('Araneides  des  lies  de  la  Reunion,'  pi.  vi.  fig-s.  1  and 
2)  gives  a  g-ood  instance  of  the  small  size  of  the  male,  in  Epeira  nigra. 
In  this  species,  as  I  may  add,  the  male  is  testaceous  and  the  female 
black  with  leg's  banded  v/ith  red.  Other  even  more  striking  cases  of 
inequality  in  size  between  the  sexes  have  been  recorded  ('Quarterly 
Journal  of  Science,*  1868,  July,  p.  429);  but  I  have  not  seen  the  origi- 
nal accounts. 


270  THE  DESCENT  OF  MAN. 

"voured,  a  sight  which,  as  he  adds,  filled  him  with  horror  and 
indignation.""'  The  Rev.  O.  P.  Cambridge"  accounts  in  the  fol- 
lowing manner  for  the  extreme  smallness  of  the  male  in  the 
genus  Nephila.  "M.  Vinson  gives  a  graphic  account  of  the  agile 
"way  in  which  the  diminutive  male  escapes  from  the  ferocity  of 
"the  female,  by  gliding  about  and  playing  hide  and  seek  over  her 
"body  and  along  her  gigantic  limbs:  in  such  a  pursuit  it  is  evi- 
"dent  that  the  chances  of  escape  would  be  in  favor  of  the  smallest 
"males  while  the  larger  ones  would  fall  early  victims;  thus  grad- 
"ually  a  diminutive  race  of  males  would  be  selected,  until  at  last 
"they  would  dwindle  to  the  smallest  possible  size  compatible  with 
"the  exercise  of  their  generative  functions, — in  fact  probably  to 
*  the  size  we  now  see  them,  i.e.,  so  small  as  to  be  a  sort  of  para- 
"site  upon  the  female,  and  either  beneath  her  notice,  or  too  agile 
"and  too  small  for  her  to  catch  without  great  difficulty." 

Westring  has  made  the  interesting  discovery  that  the  males 
ot  several  species  of  Theridion-^  have  the  power  of  making  a 
strldulating  sound,  whilst  the  females  are  mute.  The  apparatus 
consists  of  a  serrated  ridge  at  the  base  of  the  abdomen,  against 
which  the  hard  hinder  part  of  the  thorax  is  rubbed;  and  of  this 
structure  not  a  trace  can  be  detected  in  the  females.  It  deserves 
notice  that  several  writers,  including  the  well-known  arachnolo- 
gist  Walckenaer,  have  declared  that  spiders  are  attracted  by 
music.^*  From  the  analogy  of  the  Orthoptera  and  Homoptera, 
to  be  described  in  the  next  chapter,  v/e  may  feel  almost  sure 
that  the  stridulation  serves  as  Westring  also  believes,  to  call  or  to 
excite  the  female;  and  this  is  the  first  case  known  to  me  in  the  as- 
cending scale  of  the  animal  kingdom  of  sounds  emitted  for  this 
purpose.^"* 

Class,  Myriapoda. — In  neither  of  the  two  orders  in  this  class, 
the  millipedes  and  centipedes,  can  I  find  any  well-marked  in- 
stances of  such  sexual  differences  as  more  particularly  concern 
us.  In  Glomeris  limbata,  however,  and  perhaps  in  some  few 
other  species,  the  males  differ  slightly  in  color  from  the  females; 
but  this  Glomeris  is  a  highly  variable  species.    In  the  males  of 

21  Kirby  and  Spence,  'Introduction  to  Entomology,'  vol.  i.  1818,  p.  280. 

22  'Proc.  Zoolog.  Soc.,'  1871,  p.  621. 

23  Theridion  (Asagena,  Sund.)  serratipes,  4-punctatum  et  guttatum; 
see  Westring,  in  Kroyer,  'Naturhist.  Tidskrift,'  vol.  iv.  1842-1843,  p.  349; 
and  vol.  ii.  1846-1849,  p.  342.  See,  also,  for  other  species,  'Araneae 
Buecicae,'  p.  184. 

2*  Dr.  H.  H.  van  Zouteveen,  in  his  Dutch  translation  of  this  work 
(vol.  i.  p.  444),  has  collected  several  cases. 

25  Hilgendorf,  however,  has  lately  called  attention  to  an  analogous 
structure  in  some  of  the  higher  crustaceans,  which  seems  adapted  to 
produce  sound;    see  'Zoological  Record,'  1869,  p.  603. 


SPIDERS.  271 

the  Diplopoda,  the  legs  belonging  either  to  one  of  the  anterior  or 
of  the  posterior  segments  of  the  body  are  modified  into  prehensile 
hooks  which  serve  to  secure  the  female.  In  some  species  of  lulus 
the  tarsi  of  the  male  are  furnished  with  membranous  suckers  for 
the  same  purpose.  As  we  shall  see  when  we  treat  of  Insects,  it  is 
a  much  more  unusual  circumstance,  that  it  is  the  female  in  Litho- 
bius,  which  is  furnished  with  prehensile  appendages  at  the  ex- 
tremity of  her  body  for  holding  the  male.-^ 

28  Walckenaer    et    P.    Gervais,    'Hist.    Nat.    des    Insectes:   Apteres,' 
torn.  iv.  1847,  pp.  17,  19,  68. 


372  THE  DESCENT  OF  MAN. 


CHAPTER    X. 

SECONDARY   SEXUAL.  CHARACTERS   OF   INSECTS. 

Diversified  structures  possessed  by  the  males  for  seizing  the  females- 
Differences  between  the  sexes,  of  which  the  meaning  is  not  under- 
stood— Difference  in  size  between  the  sexes — Thysanura— Diptera 
— Hemiptera  — Homoptera,  musical  powers  possessed  by  the  males 
alone — Orthoptera,  musical  instruments  of  the  males,  much  diver- 
sified in  structure;  pugnacity;  colors— Neuroptera,  sexual  differ- 
ences in  color— Hymenoptera,  pugnacity  and  colors— Coleoptera, 
colors;  furnished  with  great  horns,  apparently  as  an  ornament; 
battles;  stridulating  organs  generally  common  to  both  sexes. 

In  the  immense  class  of  insects  the  sexes  sometimes  differ  in 
their  locomotive-organs,  and  often  in  their  sense-organs,  as  in 
the  pectinated  and  beautifully  plumose  antennae  of  the  males  of 
many  species.  In  Chloeon,  one  of  the  Ephemera,  the  male  has 
great  pillared  eyes,  of  which  the  female  is  entirely  destitute.^ 
The  ocelli  are  absent  in  the  females  of  certain  insects,  as  in  the 
Mutillidse;  and  here  the  females  are  likewise  wingless.  But  we 
are  chiefly  concerned  with  structures  by  which  one  male  is  en- 
abled to  conquer  another,  either  in  battle  or  courtship,  through 
his  strength,  pugnacity,  ornaments,  or  music.  The  innumerable 
contrivances,  therefore,  by  which  the  male  is  able  to  seize  the  fe- 
male, may  be  briefly  passed  over.  Besides  the  complex  struct- 
ures at  the  apex  of  the  abdomen,  which  ought  perhaps  to  be 
ranked  as   primary  organs,^   "it  is   astonishing,"   as   Mr.   B.  D. 

1  Sir  J.  Lubbock,  'Transact.  Linnean  Soc.,'  vol.  xxv.  1866,  p.  484. 
With  respect  to  the  Mutillidae,  see  Westwood,  'Modern  Class,  of  In- 
sects,' vol.  ii.  p.  213. 

2  These  organs  in  the  male  often  differ  in  clo-sely-allied  species,  and 
afford  excellent  specific  characters.  But  their  importance,  from  a 
functional  point  of  view,  as  Mr.  R.  MacLachlan  has  remarked  to  me, 
has  probably  been  overrated.  It  has  been  suggested,  that  slight  dif- 
ferences in  these  organs  would  suffice  to  prevent  the  intercrossing  of 
well-marked  varieties  or  incipient  species,  and  would  thus  aid  in 
their  development.  That  this  can  hardly  be  the  case,  we  may  infer 
from  the  many  recorded  cases  (see,  for  instance,  Bronn,  'Geschichte 
der  Natur,'   B.    ii.   1843,    s.   164;     and  Westwood,   'Transact.   Ent.   Soc.,* 


INSECTS.  273 

Walsh^  has  remarked,  "how  many  different  organs  are  worked 
"in  by  nature  for  the  seemingly  insignificant  object  of  enabling 
"the  male  to  grasp  the  female  firmly."  The  mandibles  or  jaws 
are  sometimes  used  for  this  purpose;  thus  the  male  Corydalis 
cornutus  (a  neuropterous  insect  in  some  degree  allied  to  the 
Dragon-flies,  &c.)  has  immense  curved  jaws,  many  times  longer 
than  those  of  the  female;  and  they  are  smooth  instead  of  being 
toothed,  so  that  he  is  thus  enabled  to  seize  her  without  injury.* 
One  of  the  stag-^3eetles  of  North  America  (Lucanus  elaphus)  uses 
his  jaws,  which  are  much  larger  than  those  of  the  female,  for  the 
same  purpose,  but  probably  likewise  for  fighting.  In  one  of  the 
sand-wasps  (Ammophila)  the  jaws  in  the  two  sexes  are  closely 
alike,  but  are  used  for  widely  different  purposes:  the  males,  as 
Professor  Westwood  observes,  "are  exceedingly  ardent,  seizing 
"their  partners  round  the  neck  with  their  sickle-shaped  jaws;"^ 
whilst  the  females  use  these  organs  for  burrowing  in  sand-banks 
and  making  their  nests. 

The  tarsi  of  the  front-legs  are  dilated  in  many  male  beetles,  or 
are  furnished  with  broad  cushions  of  hairs;  and  in  many  genera 
of  water-beetles  they  are  armed  with  a  round  flat  sucker,  so  that 
the  male  may  adhere  to  the  slippery  body  of  the  female. 
It  is  a  much  more  unusual  circumstance  that  the  female 
of  some  water-beetles  (Dytiscus)  have  their  elytra  deeply 
grooved,  and  in  Acilius  sulcatus  thickly  set  with  hairs, 
as  an  aid  to  the  male.  The  females  of  some  other 
water-beetles  (Hydroporus)  have  their  elytra  punctured  for 
the  same  purpose.^  In  the  male  of  Crabro  cribrarius  (fig. 
9),  it  is  the  tibia  which  is  dilated  into  a  broad  horny  plate, 
with  minute  membranous  dots,  giving  to  it  a  singular  appear- 
ance like  that  of  a  riddle.^    In  the  male  of  Penthe  (a  genus  of 

vol.  iii.  1842,  p.  195)  of  distinct  species  having-  been  observed  in  union. 
Mr.  MacLacMan  informs  me  (vide  'Stett.  Ent.  Zeitung,'  1867,  s.  155) 
that  when  several  species  of  Phryganidae,  which  present  strongly- 
pronounced  differences  of  this  kind,  were  confined  together  by  Dr. 
Aug.  Meyer,  they  coupled,  and  one  pair  produced  fertile  ova. 

3  'The  Practical  Entomologist,  Philadelphia,  vol.  ii.  May,  1867,  p.  88. 

*Mr.  Walsh,  ibid.   p.   107. 

5  'Modern  classification  of  Insects,'  vol.  ii.  1840,  pp.  205,  206.  Mr. 
Walsh,  who  called  my  attention  to  the  double  use  of  the  jaws,  says 
that  he  has  repeatedly  observed  this  fact. 

®  We  have  here  a  curious  and  inexplicable  case  of  dimorphism,  for 
some  of  the  females  of  four  European  species  of  Dytiscus,  and  of 
certain  species  of  Hydroporus,  have  their  elytra  smooth;  and  no  in- 
termediate gradations  between  the  sulcated  or  punctured,  and  the 
quite  smooth  elytra  have  been  observed.  See  Dr.  H.  Schaum,  as 
quoted  in  the  'Zoologist,'  vol.  v.-vi.  1847-48,  p.  1896.  Also  Kirby  and 
Spence,  'Introduction  to  Entomology,'  vol.  iii.  1826,  p.  305. 

7  Westwood,  'Modern  Class,  vol.  ii.  p.  193.  The  following  statement 
19 


274 


THE  DESCENT  OF  MAN. 


beetles)  a  few  of  the  middle  joints  of  the  antennae  are  dilated  and 
furnished  on  the  inferior  surface  with  cushions  of  hair,  exactly 

like  those  on  the  tarsi  of  the  Car- 
abidaB,  "and  obviously  for  the  same 
end."  In  male  dragon-flies,  "the 
"appendages  at  the  tip  of  the  tail 
"are  modified  in  an  almost  infinite 
"variety  of  curious  patterns  to  en- 
"able  them  to  embrace  the  neck  of 
"the  female."  Lastly,  in  the  males 
of  many  insects,  the  legs  are  fur- 
nished with  peculiar  spines,  knobs 
or  spurs;  or  the  whole  leg  is  bowed 
or  thickened,  but  this  is  by  no 
means  invariably  a  sexual  charac- 
ter; or  one  pair,  or  all  three  pairs 
are  elongated,  sometimes  to  an  ex- 
travagant length.^ 

The  sexes  of  many  species  in  all 
the  orders  present  differences,  of 
which  th©  meaning  is  not  under- 
stood. One  curious  case  is  that  of  a 
beetle  (fig.  10),  the  male  of  which 
has  the  left  mandible  much  en- 
larged; so  that  the  mouth  is  greatly 
distorted.  In  another  Carabidous  beetle,  Eurygnathus,^  we  have 
the  case,  unique  as  far  as  known  to  Mr.  Wollaston,  of  the  head  of 
the  female  being  much  broader  and  larger,  though  in  a  variable  de- 
gree, than  that  of  the  male.  Any  number  of  such  cases  could  be 
given.  They  abound  in  the  Lepidoptera:  one  of  the  most  extra- 
ordinary is  that  certain  male  butterflies  have  their  fore-legs  more 
or  less  atrophied,  with  the  tibiae  and  tarsi  reduced  to  mere  rudi- 
mentary knobs.  The  wings,  also,  in  the  two  sexes  often  differ  in 
neuration,^*'  and  sometimes  considerably  in  outline,  as  in  the 
Aricoris  epitus,  which  v/as  shown  to  me  in  the  British  Museum 
by  Mr.  A.  Butler.  The  males  of  certain  South  American  butter- 
flies have  tufts  of  hair  on  the  margins  of  the  wings,  and  horny 


Fig.  9.  Crabro  cribrarius.  Up- 
per figure,  male;  lower  fig- 
ure, female. 


about  Penthe,  and  others  in  inverted  commas,  are  taken  from  Mr. 
Walsh,   'Practical  Entomologist,'   Philadelphia,   vol.   ii.   p.  88. 

8  Kirby  and  Spence,   'Introduct.'  &c.,  vol.  iii.  pp.  332-336. 

»  'Insecta  Maderensia,'  1854,  p.  20. 

10  E.  Doubleday,  'Annals  and  Mag.  of  Nat.  Hist.'  vol.  i.  1848,  p.  379. 
I  may  add  that  the  wings  in  certain  Hymenoptera  (see  Shuckard, 
'Fossorial  Hymenop.'  1837,  pp.  39-43)  differ  in  neuration  according  to 
sex. 


INSECTS. 


275 


excrescences  on  the  discs  of  the  posterior  pair."  In  several  Brit- 
ish butterflies,  as  shown  by  Mr.  Wonfor,  the  males  alone  are  in 
parts  clothed  with  peculiar  scales. 

The  use  of  the  bright  light  of  the  female  glow- 
worm has  been  subject  to  much  discussion.  The 
male  is  feebly  luminous,  as  are  the  larvae  and 
even  the  eggs.  It  has  been  supposed  by  some 
authors  that  the  light  serves  to  frighten  away 
enemies,  and  by  others  to  guide  the  male  to  the 
female.  At  last,  Mr.  Belt"'  appears  to  have 
solved  the  difficulty:  he  finds  that  all  the 
Lampyridse  which  he  has  tried  are  highly  dis- 
tasteful to  insectivorous  mammals  and  birds. 
Hence  it  is  in  accordance  with  Mr.  Bates'  view, 
hereafter  to  be  explained,  that  many  insects 
mimic  the  Lampyridse  closely,  in  order  to  be 
mistaken  for  them,  and  thus  to  escape  destruc- 
tion. He  further  believes  that  the  luminous 
species  profit  by  being  at  once  recognized  as  un- 
palatable. It  is  probable  that  the  same  explana- 
tion may  be  extended  to  the  Elaters,  both  sexes 
of  which  are  highly  luminous.  It  is  not  known 
why  the  wings  of  the  female  glow-worm  have 
not  been  developed;  but  in  her  present  state  she 
closely  resembles  a  larva,  and  as  larvae  are  so 
largely  preyed  on  by  many  animals,  we  can  un- 
derstand why  she  has  been  rendered  so  much 
more  luminous  and  conspicuous  than  the  male; 
and  why  the  larvae  themselves  are  likewise 
luminous. 


Difference  in  Size  between  the  Sexes.  —With 
insects  of  all  kinds  the  males  are  commonly 
smaller  than  the  females;  and  this  difference 
can  often  be  detected  even  in  the  larval  state. 
So  considerable  is  the  difference  between  the 
male  and  female  cocoons  of  the  silk-moth  (Bom- 
byx  mori),  that  in  France  they  are  separated 
by  a  particular  mode  of  weighing.^^  In  the  lower  classes  of  the 
animal  kingdom,  the  greater  size  of  the  females  seems  generally 


Fig-.  10.  Taphro- 
deres  distortus 
(much  enlarg- 
ed). Upper  fig-- 
u  r  e,  male; 
lower  figure, 
female. 


"H.  W.  Bates,  in  'Journal  of  Proc.  Linn.  Soc'  vol.  vi.  1862,  p.  74. 
Mr.  Wonfor's  observations  are  quoted  in  'Popular  Science  Review,' 
1868,  p.  343. 

12  'The  Naturalist  in  Nicaragua,'  1874,  pp.  316-320.  On  the  phosphor- 
escence of  the  eggs,  see  'Annals  and  Mag.  of  Nat.  Hist.'  1871,  Nov., 
p.  372. 

isRobinet,   'Vers  a  Sole,'  1848,  p.  207. 


276  THE  DESCENT  OF  MAN. 

to  depend  on  their  developing  an  enormous  number  of  ova;  and 
this  may  to  a  certain  extent  hold  good  with  insects.  But  Dr. 
Wallace  has  suggested  a  much  more  probable  explanation.  He 
finds,  after  carefully  attending  to  the  development  of  the  cater- 
pillars of  Bombyx  cynthia  and  yamamai,  and  especially  to  that 
of  some  dwarfed  caterpillars  reared  from  a  second  brood  on  un- 
natural food,  "that  in  proportion  as  the  individual  moth  is  finer, 
"so  is  the  time  required  for  its  metamorphosis  longer;  and  for 
"this  reason  the  female,  which  is  the  larger  and  heavier  insect, 
"from  having  to  carry  her  numerous  eggs,  will  be  preceded  by  the 
"male,  which  is  smaller  and  has  less  to  mature.""  Now  as  most 
insects  are  short-lived,  and  as  they  are  exposed  to  many  dangers, 
it  would  manifestly  be  advantageous  to  the  female  to  be  impreg- 
nated as  soon  as  possible.  This  end  would  be  gained  by  the 
males  being  first  matured  in  large  numbers  ready  for  the  advent 
of  the  females;  and  this  again  would  naturally  follow,  as  Mr.  A. 
R.  Wallace  has  remarked,^^  through  natural  selection;  for  the 
smaller  males  would  be  first  matured,  and  thus  would  procreate 
a  large  number  of  offspring  which  would  inherit  the  reduced  size 
of  their  male  parents,  whilst  the  larger  males  from  being  ma- 
tured later  would  leave  fev/er  offspring. 

There  are,  however,  exceptions  to  the  rule  of  male  insects  be- 
ing smaller  than  the  females:  and  some  of  these  exceptions  are 
intelligible.  Size  and  strength  would  be  an  advantage  to  the 
males,  which  fight  for  the  possession  of  the  females;  and  in  these 
cases,  as  v/ith  the  stag-beetle  (Lucanus),  the  males  are  larger 
than  the  females.  There  are,  however,  other  beetles  which  are  not 
known  to  fight  together,  of  which  the  males  exceed  the  females 
in  size;  and  the  meaning  of  this  fact  is  not  known;  but  in  some 
of  these  cases,  as  with  the  huge  Dynastes  and  Megasoma,  we  can 
at  least  see  that  there  would  be  no  necessity  for  the  males  to  be 
smaller  than  the  females,  in  order  to  be  matured  before  them,  for 
these  beetles  are  not  short-lived,  and  there  would  be  ample  time  for 
the  pairing  of  the  sexes.  So  again,  male  dragon-fiies  (Libellu- 
lidse)  are  sometimes  sensibly  larger,  and  never  smaller,  than  the 
females;"  and  as  Mr.  MacLachlan  believes,  they  do  not  generally 
pair  with  the  females  until  a  week  or  fortnight  has  elapsed,  and 
until  they  have  assumed  their  proper  masculine  colors.  But  the 
most  curious  cases,  showing  on  what  complex  and  easily  over- 
looked relations,  so  trifling  a  character  as  difference  in  size  be- 
tween the  sexes  may  depend,  is  that  of  the  aculeate  Hymenop- 


"  'Transact.  Ent.  Soc'  3rd  series,  voL  v.  p.  486. 

IB  'Journal  of  Proc.  Ent.  Soc'  Feb.  4th,  1867,  p.  Ixxi. 

16  For  this  and  other  statements  on  the  size  of  the  sexes,  see  Kirby 
and  Spence,  ibid.  vol.  iii.  p.  300;  on  the  duration  of  life  in  insects, 
see  p.  344. 


^  THYSANURA.  27^ 

tera;  for  Mr.  F.  Smith  informs  me  that  throughout  nearly  the 
whole  of  this  large  group,  the  males,  in  accordance  with  the  gen- 
eral rule,  are  smaller  than  the  females,  and  emerge  about  a  week 
before  them;  but  amongst  the  Bees,  the  males  of  Apis  mellifica, 
Anthidium  manicatum,  and  Anthophora  acervorum,  and  amongst 
the  Fossores,  the  males  of  the  Methoca  ichneumonides,  are  larger 
than  the  females.  The  explanation  of  this  anomaly  is  that  a 
marriage  flight  is  absolutely  necessary  with  these  species,  and  the 
male  requires  great  strength  and  size  in  order  to  carry  the  female 
through  the  air.  Increased  size  has  here  been  acquired  in  oppo- 
sition to  the  usual  relation  between  size  and  the  period  of  de- 
velopment, for  the  males,  though  larger,  emerge  before  tne 
smaller  females. 

We  will  now  review  the  several  Orders,  selecting  such  facts  as 
more  particularly  concern  us.  The  Lepidoptera  (Butterflies  and 
Moths)  will  be  retained  for  a  separate  chapter. 

Order,  Tliysanura.— The  members  of  this  lowly  organized  order 
are  wingless,  dull-colored,  minute  insects,  with  ugly,  almost 
misshapen  heads  and  bodies.  Their  sexes  do  not  differ;  but  they 
are  interesting  as  showing  us  that  the  males  pay  sedulous  court 
to  the  females  even  low  down  in  the  animal  scale.  Sir  J.  Lub- 
bock^' says:  "it  is  very  amusing  to  see  these  little  creatures 
"(Smynthurus  luteus)  coquetting  together.  The  male,  which  is 
"much  smaller  than  the  female,  runs  round  her,  and  they  butt 
"one  another,  standing  face  to  face,  and  moving  backward  and 
"forward  like  two  playful  lambs.  Then  the  female  pretends  to 
"run  away  and  the  male  runs  after  her  with  a  queer  appearance 
"of  anger,  gets  in  front  and  stands  facing  her  again;  then  she 
"turns  coyly  round,  but  he,  quicker  and  more  active,  scuttles 
"round  too,  and  seems  to  whip  her  with  his  antennae;  then  for  a 
"bit  they  stand  face  to  face,  play  with  their  antennae,  and  seem 
"to  be  all  in  all  to  one  another." 

Order,  Diptera  (Flies).— The  sexes  differ  little  in  color.  The 
greatest  difference,  known  to  Mr.  F.  Walker,  is  in  the  genus 
Bibio,  in  which  the  males  are  blackish  or  quite  black,  and  the 
females  obscure  brownish-orange.  The  genus  Elaphomyia,  dis- 
covered by  Mr.  Wallace^^  in  New  Guinea,  is  highly  remarkable, 
as  the  males  are  furnished  with  horns,  of  which  the  females  are 
quite  destitute.  The  horns  spring  from  beneath  the  eyes,  and 
curiously  resemble  those  of  a  stag,  being  either  branched  or  pal- 
mated.  In  one  of  the  species,  they  equal  the  whole  body  in 
length.  They  might  be  thought  to  be  adapted  for  fighting,  but 
as  in  one  species  they  are  of  a  beautiful  pink  color,  edged  with 

"'Transact.   Linnean  Soc'  vol.   xxvi.  1868,  p.  296. 
18  'The  Malay  Archipelago,'  vol.  iL  1869,  p.  313. 
19 


278  THE   DESCENT  OF  MAN. 

black,  with  a  pale  central  stripe,  and  as  these  insects  have  alto- 
gether a  very  elegant  appearance,  it  is  perhaps  more  probable 
that  they  serve  as  ornaments.    That  the  males  of  some  Diptera 
fight  together  is  certain;  for  Prof.  Westwood^'  has  several  times 
seen  this  with  the  Tipulae.    The  males  of  other  Diptera  appar- 
ently try  to  win  the  females  by  their  music:  H.  Muller-o  watched 
for  some  time  two  males  of  an  Eristalis  courting  a  female;  they 
hovered  above  her,  and  flew  from  side  to  side,  making  a  high 
humming  noise  at  the  same  time.    Gnats  and  mosquitoes  (Culi- 
cidse)   also  seem  to  attract  each  other  by  humming;  and  Prof. 
Mayer  has  recently  ascertained  that  the  hairs  on  the  antennae  of 
the  male  vibrate  in  unison  with  the  notes  of  a  tuning-fork,  within 
the  range  of  the  sounds   emitted   by  the   female.    The   longer 
hairs  vibrate  sympathetically  with  the  graver   notes,   and  the 
shorter  hairs  with  the  higher  ones.    Landois  also  asserts  that  he 
has  repeatedly  drawn  down  a  whole  swarm  of  gnats  by  uttering 
a  particular  note.    It  may  be  added  that  the  mental  faculties  of 
the  Diptera  are  probably  higher  than  in  most  other  insects,  in  ac- 
cordance with  their  highly  developed  nervous  system.^^ 

Order,  Hemiptera  (Field-Bugs).— Mr.  J.  W.  Douglas,  who  has 
particularly  attended  to  the  British  species,  has  kindly  given  me 
an  account  of  their  sexual  differences.  The  males  of  some  species 
are  furnished  with  wings,  whilst  the  females  are  wingless;  the 
sexes  differ  in  the  form  of  their  bodies,  elytra,  antennae  and  tarsi; 
but  as  the  signification  of  these  differences  are  unknown  they 
may  be  here  passed  over.  The  females  are  generally  larger  and 
more  robust  than  the  males.  With  British,  and,  as  far  as  Mr. 
Douglas  knows,  with  exotic  species,  the  sexes  do  not  commonly 
differ  much  in  color;  but  in  about  six  British  species  the  male  is 
considerably  darker  than  the  female,  and  in  about  four  other 
species  the  female  is  darker  than  the  male.  Both  sexes  of  some 
species  are  beautifully  colored;  and  as  these  insects  emit  an 
extremely  nauseous  odor,  their  conspicuous  colors  may  serve  as 
a  signal  that  they  are  unpalatable  to  insectivorous  animals.  In 
some  few  case  their  colors  appear  to  be  directly  protective:  thus 
Prof.  Hoffmann  informs  me  that  he  could  hardly  distinguish  a 
small  pink  and  green  species  from  the  buds  on  the  trunks  of  lime- 
trees,  which  this  insect  frequents. 

Some  species  of  Reduvidae  makes  a  stridulating  noise;    and,  in 


19  'Modern  Classification  of  Insects,'  vol.   ii.  1840,  p.  526, 

20  Anwendung,  &c.,  'Verb.  d.  n.  V.  Jahrg-.*  xxix.  p.  80.  Mayer,  in 
'American  Naturalist,'  1874,  p.  236. 

21  See  Mr.  B.  T.  Lowne's  interesting  work,  'On  the  Anatomy  of  the 
Blow-fly,  Musca  vomitoria,'  1870,  p.  14.  He  remarks  (p.  33)  that,  "the 
"captured  flies  utter  a  peculiar  plaintive  note,  and  that  this  sound 
"causes  other  flies  to  disappear." 


HEMIPTERA   AND    HOMOPTERA.  279 

the  case  of  Pirates  stridulus,  this  is  said^^  to  be  effected  by  the 
movement  of  the  neck  within  the  pro-thoracic  cavity.  Accord- 
ing to  Westring,  Reduvius  personatus  also  stridulates.  But  I 
have  no  reason  to  suppose  that  this  is  a  sexual  character,  except- 
ing that  with  non-social  insects  there  seems  to  be  no  use  for 
sound-producing  organs,  unless  it  be  as  a  sexual  call. 

Order,  Homoptera. — Every  one  who  has  wandered  in  a  tropical 
forest  must  have  been  astonished  at  the  din  made  by  the  male 
Cicadae.  The  females  are  mute;  as  the  Grecian  poet  Xenarchus 
says,  "Happy  the  Cicadas  live,  since  they  all  have  voiceless 
"wives."  The  noise  thus  made  could  be  plainly  heard  on  board 
the  "Beagle,"  when  anchored  at  a  quarter  of  a  mile  from  the 
shore  of  Brazil;  and  Captain  Hancock  says  it  can  be  heard  at  the 
distance  of  a  mile.  The  Greeks  formerly  kept,  and  the  Chinese 
now  keep  these  insects  in  cages  for  the  sake  of  their  song,  so  that 
it  must  be  pleasing  to  the  ears  of  some  men.-^'  The  Cicadidse 
usually  sing  during  the  day,  whilst  the  Pulgoridae  appear  to  be 
night-songsters.  The  sound,  according  to  Landois,'*  is  produced 
by  the  vibration  of  the  lips  of  the  spiracles,  which  are  set  into 
motion  by  a  current  of  air  emitted  from  the  tracheae  but  this  view 
has  lately  been  disputed.  Dr.  Powell  appears  to  have  proved-^ 
that  it  is  produced  by  the  vibration  of  a  membrane,  set  into  ac- 
tion by  a  special  muscle.  In  the  living  insect,  whilst  stridulat- 
ing,  this  membrane  can  be  seen  to  vibrate;  and  in  the  dead  in- 
sect the  proper  sound  is  heard,  if  the  muscle,  when  a  little  dried 
and  hardened,  is  pulled  with  the  point  of  a  pin.  In  the  female 
the  whole  complex  musical  apparatus  is  present,  but  is  much  less 
developed  than  in  the  male,  and  is  never  used  for  producing 
sound. 

With  respect  to  the  object  of  the  music.  Dr.  Hartman,  in 
speaking  of  the  Cicada  septemdecim  of  the  United  States,  says,-® 
"the  drums  are  now  (June  6th  and  7th,  1851)  heard  in  all  di- 
"rections.  This  I  believe  to  be  the  marital  summons  from  the 
"males.  Standing  in  thick  chestnut  sprouts  about  as  high  as  my 
"head,  where  hundreds  were  around  me,  I  observed  the  females 
"coming  around  the  drumming  males."  He  adds,  "this  season 
"(Aug.  1868)  a  dwarf  pear-tree  in  my  garden  produced  about  fifty 
"larvae  of  Cic.  pruinosa;  and  I  several  times  noticed  the  females 

22  Westwood,   'Modern  Class,  of  Insects,'  vol.  ii.  p.  473. 

23  These  particulars  are  tsiken  from  Westwood's  'Modern  Class,  of 
Insects,'  vol.  ii.  1840,  p.  422.  See,  also,  on  the  Fulgoridae,  Kirby  and 
Spence,    'Introduct.'  vol.   ii.   p.  401. 

2*  'Zeitschrift  fur  wissenschaft  Zoolog-.'   B.   xvii.  1867,   s.  152-158. 
25  'Transact.  New  Zealand  Institute,'  vol.  v.  1873,  p.  286. 
28 1  am  indebted  to  Mr.  Walsh  for  having  sent  me  this  extract  from 
a  'Journal  of  the  Doings  of  Cicada  septemdecim,'  by  Dr.  Hartman. 


2g0  THE  DESCENT  OF  MAN. 

"to  alight  near  a  male  while  he  was  uttering  his  clanging  notes." 
Fritz  Miiller  writes  to  me  from  S.  Brazil  that  he  has  often  lis- 
tened to  a  musical  contest  between  two  or  three  males  of  a  species 
with  a  particularly  loud  voice,  seated  at  a  considerable  distance 
from  each  other:  as  soon  as  one  had  finished  his  song,  another 
immediately  began,  and  then  another.  As  there  is  so  much  ri- 
valry between  the  males,  it  is  probable  that  the  females  not  only 
find  them  by  their  sounds,  but  that,  like  female  birds,  they  are 
excited  or  allured  by  the  male  with  the  most  attractive  voice. 

I  have  not  heard  of  any  well-marked  cases  of  ornamental  dif- 
ferences between  the  sexes  of  the  Homoptera.  Mr.  Douglas  in- 
forms me  that  there  are  three  British  species,  in  which  the  male 
is  black  or  marked  with  black  bands,  whilst  the  females  are  pale- 
colored  or  obscure. 

Order,  Orthoptera  (Crickets  and  Grasshoppers). — The  males  in 
the  three  saltatorial  families  in  this  Order  are  remarkable  for 
their  musical  powers,  namely  the  Achetidse  or  crickets,  the 
LocustidsB  for  which  there  is  no  equivalent  English  name,  and 
the  Acridiidae  or  grasshoppers.  The  stridulation  produced  by 
some  of  the  Locustidse  is  so  loud  that  it  can  be  heard  during  the 
night  at  the  distance  of  a  mile;^^  and  that  made  by  certain  species 
is  not  unmusical  even  to  the  human  ear,  so  that  the  Indians  on  the 
Amazons  keep  them  in  wicker  cages.  All  observers  agree  that 
the  sounds  serve  either  to  call  or  excite  the  mute  females.  With 
respect  to  the  migratory  locusts  of  Russia,  Korte  has  given^  an 
interesting  case  of  selection  by  the  female  of  a  male.  The  males 
of  this  species  (Pachytylus  migratorius)  whilst  coupled  with  the 
female  stridulate  from  anger  or  jealousy,  if  approached  by  other 
males.  The  house-cricket  when  surprised  at  night  uses  its  voice 
to  warn  its  fellows.-®  In  North  America  the  Katy-did  (Platy- 
phyllum  concavum,  one  of  the  Locustidse)  is  described^  as  mount- 
ing on  the  upper  branches  of  a  tree,  and  in  the  evening  beginning 
"'his  noisy  babble,  while  rival  notes  issue  from  the  neighboring 
"trees,  and  the  groves  resound  with  the  call  of  Katy-did-she-did 
"the  live-long  night."  Mr.  Bates,  in  speaking  of  the  European 
field-cricket  (one  of  the  Achetidse),  says,  "the  male  has  been  ob- 
"served  to  place  himself  in  the  evening  at  the  entrance  of  his 
"burrow,  and  stridulate  until  a  female  approaches,  when  the 
"louder  notes  are  succeeded  by  a  more  subdued  tone,  whilst  the 

27  L.  Guilding,   'Transact  Linn.  Soc'  vol,  xv.  p.  154. 

2^  I  state  this  on  the  authority  of  Koppen,  'Ueber  die  Heuschrecken 
in  Sudrussland,'  1866,  p.  32,  for  I  have  in  vain  endeavored  to  procure 
Korte' s  work. 

2»  Gilbert  White,  'Nat.  Hist,  of  Selborne,'  vol.  ii.  1825,  p.  262. 

*»  Harris,  'Insects  of  New  England,'  1842,  p.  128. 


ORTHOPTERA. 


281 


"successful  musician  caresses  -with  his  antennae  the  mate  he  has 
"won."^^  Dr.  Scudder  was  able  to  excite  one  of  these  insects  to 
answer  him  by  rubbing  on  a  file  with  a  quill.®-  In  both  sexes  a  re- 
markable auditory  apparatus  has  been  discovered  by  Von  Siebold, 
situated  in  the  front  legs.^^ 

In  the  three  Families  the  sounds  are  differently  produced.  In 
the  males  of  the  Achetidae  both  wing-covers  have  the  same  ap- 
paratus; and  this  in  the  field-cricket  (Gryllus  campestris,  fig.  11) 
consists,  as  described  by  Landois,^*  of  from  131  to  138  sharp,  trans- 
verse ridges  or  teeth  (st)  on  the  under  side  of  one  of  the  nervures 


Fig-.      11.      Gryllus   campestris    (from   Landois). 

Rig-ht-hand  figure,  under  side    of    part    of    a 

wing-nervure,  much  magnified,    showing    the 

teeth,  St. 
Left-hand  figure,  upper  surface  of  wing-cover, 

with  the  projecting,  smooth  nervure  r,  across 

which  the   teeth   (st)    are  scraped. 


Fig.  12.  Teeth  of 
Nervure  of  Gryl- 
lus domesticus 
(from    Landois). 


of  the  wing-cover.  This  toothed  nervure  is  rapidly  scraped  across 
a  projecting,  smooth,  hard  nervure  (r)  on  the  upper  surface  of 
the  opposite  wing.  First  one  wing  is  rubbed  over  the  other,  and 
then  the  movement  is  reversed.  Both  wings  are  raised  a  little 
at  the  same  time,  so  as  to  increase  the  resonance.  In  some  spe- 
cies the  wing-covers  of  the  males  are  furnished  at  the  base  with 
a  talc-like  plate.^"    I  here  give  a  drawing  (fig.  12)  of  the  teeth 


31  'The  Naturalist  on  the  Amazons,'  vol.  i.  1863,  p.  252.  Mr,  Bates 
gives  a  very  interesting  discussion  on  the  gradations  in  the  musical 
apparatus  of  the  three  families.  See,  also,  Westwood,  'Modern 
Class.*  vol.  ii.  pp.  445  and  453. 

32  'Proc.  Boston  Soc.  of  Nat.  Hist.'  vol.  xi.  April  1868. 

33  'Nouveau  Manuel  d'Anat.  Comp.'  (French  translat.),  torn.  i.  1850, 
p.  567. 

a*  'Zeitschrift  fur  wissenschaft.  Zoolog.'  B.  xvii.  1867,  s.  117, 
^  Westwood,   'Modern  Class,  of  Insects,'  vol.  i,  p.  440, 


282 


THE  DESCENT  OF  MAN. 


on  the  undftr  side  of  the  nervure  of  another  species  of  Gryllus, 
viz.,  G.  domeaticus.  With  respect  to  the  formation  of  these  teeth. 
Dr.  Gruber  has  ehown^^  that  they  have  been  developed  by  the 
aid  of  selection,  n^om  the  minute  scales  and  hairs  with  which  the 
wings  and  body  sn  e  covered,  and  I  came  to  the  same  conclusion 


Fig.  13.    Chlorocoelus  Tanana  (from  Bates). 

wing-covers. 


a,  b.  Lobes  oi  opposite 


with  respect  to  those  of  the  Coleoptera.  But  Dr.  Gruber  further 
shows  that  their  development  is  in  part  directly  due  to  the  stim- 
ulus from  the  friction  of  one  wing  over  the  other. 

In  the  Locustidse  the  opposite  wing-covers  differ  from  each 
other  in  structure  (fig.  13),  and  the  action  cannot,  as  in  the  last 
family,  be  reversed.  The  left  wing,  which  acts  as  the  bow,  lies 
over  the  right  wing  which  serves  as  the  fiddle.  One  of  the  ner- 
vures  (a)  on  the  under  surface  of  the  former  is  finely  serrated, 
and  is  scraped  across  the  prominent  nervures  on  the  upper  sur- 
face of  the  opposite  or  right  wing.  In  our  British  Phasgonura 
viridissima  it  appeared  to  me  that  the  serrated  nervure  is  rubbed 


36  'Ueber  der  Tonapparat  der  Locustiden,  ein  Beitrag  zum  Darwin- 
ismus,'   'Zeitsch.  fur  wissensch.  Zoolog.'  B.  xxii.  1872,  p.  100. 


ORTHOPTERA.  283 

against  the  rounded  hind-corner  of  the  opposite  wing,  the  edge 
of  which  is  thickened,  colored  brown,  and  very  sharp.  In  the 
right  wing,  but  not  in  the  left,  there  is  a  little  plate,  as  transpar- 
ent as  talc,  surrounded  by  nervures,  and  called  the  speculum.  In 
Ephippiger  vitium,  a  member  of  the  same  family,  we  have  a 
curious  subordinate  modification;  for  the  wing-covers  are  great- 
ly reduced  in  size,  but  "the  posterior  part  of  the  pro-thorax  is 
"elevated  into  a  kind  of  dome  over  the  wing-covers,  and  which 
"has  probably  the  effect  of  increasing  the  sound. "^' 

We  thus  see  that  the  musical  apparatus  is  more  differentiated 
or  specialized  in  the  Locustidse  (which  include,  I  believe,  the 
most  powerful  performers  in  the  Order),  than  in  the  Achetidse,  in 
which  both  wing-covers  have  the  same  structure  and  the  same 
function.'*^  Landois,  however,  detected  in  one  of  the  Locustidae, 
namely  in  Decticus,  a  short  and  narrow  row  of  small  teeth,  mere 
rudiments,  on  the  inferior  surface  of  the  right  wing-cover,  which 
underlies  the  other  and  is  never  used  as  the  bow.  I  observed  the 
same  rudimentary  structure  on  the  under  side  of  the  right  wing- 
cover  in  Phasgonura  viridissima.  Hence  we  may  infer  with  con- 
fidence that  the  Locustidae  are  descended  from  a  form,  in  which, 
as  in  the  existing  Achetidae,  both  wing-covers  had  serrated  ner- 
vures on  the  under  surface,  and  could  be  indifferently  used  as  the 
bow;  but  that  in  the  Locustidae  the  two  wing-covers  gradually 
became  differentiated  and  perfected,  on  the  principle  of  the  divis- 
ion of  labor,  the  one  to  act  exclusively  as  the  bow,  and  the  other 
as  the  fiddle.  Dr.  Gruber  takes  the  same  view,  and  has  shown 
that  rudimentary  teeth  are  commonly  found  on  the  inferior  surface 
of  the  right  wing.  By  what  steps  the  more  simple  apparatus  in 
the  Achetidae  originated,  we  do  not  know,  but  it  is  probable  that 
the  basal  portions  of  the  wing-covers  originally  overlapped  each 
other  as  they  do  at  present;  and  that  the  friction  of  the  nervures 
produced  a  grating  sound,  as  is  now  the  case  with  the  wing- 
covers  of  the  females.^*  A  grating  sound  thus  occasionally  and 
accidentally  made  by  the  males,  if  it  served  them  ever  so  little  as 
a  love-call  to  the  females,  might  readily  have  been  intensified 
through  sexual  selection,  by  variations  in  the  raughness  of  the 
nervures  having  been  continually  preserved. 

In  the  last  and  third  Family,  namely  the  Aeridiidae  or  grass- 
hoppers, the  stridulation  is  produced  in  a  very  different  manner, 
and  according  to  Dr.  Scudder,  is  not  so  shrill  as  in  the  preceding 
Families.  The  inner  surface  ot  the  Femur  (fig.  14,  r)  is  fur- 
nished with  a  longitudinal  row  of  minute,  elegant,  lancet-shaped, 

s'^  Westwood,   'Modern  Class,  of  Insects,'  vol.  i.  p.  453. 

38  Landois,  'Zeltsch.  f.  wiss.  Zoolog.*  B.  xvii.  1867,  s.  121,  122. 

8»  Mr.  Walsh  also  informs  me  that  he  has  noticed  that  the  female 
of  the  Platyphyllum  concavum,  "when  captured  makes  a  feeble  grat- 
"ingr  noise  by  shuffling  her  wiiag-covers  togetheTo'' 


284 


THE  DESCENT  OF  MAN. 


Fig.  14.  Hind-leg  of  Stenoboth- 
rus  pratorum:  r,  the  stridu- 
lating  ridge;  lower  figure,  the 
teeth  forming  the  ridge,  much 
magnified  (from  Landois). 


elastic  teeth,  from  85  to  93  in  number;*"  and  these  are  scraped 
across  the  sharp,  projecting  nervures  on  the  wing-covers,  which 

are  thus  made  to  vibrate  and  re- 
sound. Harris"  says  that  when 
one  of  the  males  begins  to  play, 
he  first  "bends  the  shank  of 
"the  hind-leg  beneath  the  thigh, 
"where  it  is  lodged  in  a  furrow 
"designed  to  receive  it,  and  then 
"draws  the  leg  briskly  up  and 
"down.  He  does  not  play  both 
"fiddles  together,  but  alternately, 
"first  upon  one  and  then  on  the 
"other."  In  many  species,  the 
base  of  the  abdomen  is  hollowed 
out  into  a  great  cavity  which  is 
believed  to  act  as  a  resounding 
board.  In  Pneumora  (fig.  15),  a 
S.  African  genus  belonging  to  the 
same  family,  we  meet  with  a 
new  and  remarkable  modification; 
in  the  males  a  small  notched 
ridge  projects  obliquely  from  each  side  of  the  abdomen,  against 
which  the  hind  femora  are  rubbed.*^  As  the  male  is  furnished 
with  wings  (the  female  being  wingless),  it  is  remarkable  that  the 
thighs  are  not  rubbed  in  the  usual  manner  against  the  wing- 
covers;  but  this  may  perhaps  be  accounted  for  by  the  unusually 
small  size  of  the  hind-legs.  I  have  not  been  able  to  examine  the 
inner  surface  of  the  thighs,  which,  judging  from  analogy,  would 
be  finely  serrated.  The  species  of  Pneumora  have  been  more  pro- 
foundly modified  for  the  sake  of  stridulation  than  any  other 
orthopterous  insect;  for  in  the  male  the  whole  body  has  been 
converted  into  a  musical  instrument,  being  distended  with  air, 
like  a  great  pellucid  bladder,  so  as  to  increase  the  resonance. 
Mr.  Trimen  informs  me  that  at  the  Cape  of  Good  Hope  these 
insects  make  a  wonderful  noise  during  the  night. 

In  the  three  foregoing  families,  the  females  are  almost  always 
destitute  of  an  efiicient  musical  apparatus.  But  there  are  a  few 
exceptions  to  this  rule,  for  Dr.  Gruber  has  shown  that  both  sexes 
of  Ephippiger  vitium  are  thus  provided;  though  the  organs  differ 
in  the  male  and  female  to  a  certain  extent.  Hence  we  cannot 
suppose  that  they  have  been  transferred  from  the  male  to  the 
female,  as  appears  to  have  been  the  case  with  the  secondary  sexual 


40  Landois,  ibid.  s.  113. 

*i  'Insects  of  New  England,'  1842,  p.  133. 

*2'WeBtwood,  'Modem  Classification,'  vol. 


i.  p.  462. 


ORTHOPTERA. 


285 


characters  of  many  other  animals.  They  must  have  been  inde- 
pendently developed  in  the  two  sexes,  which  no  doubt  mutually 
call  to  each  other  during  the  season  of  love.  In  most  other  Lo- 
custidse  (but  not  according  to  Landois  in  Decticus)  the  females 


Fig.  15.    Pneumora  (from  specimens  in  the  British  Museum).    Upper 
figure,  male;  lower  figxire,  female. 

have  rudiments  of  the  stridulatory  organs  proper  to  the  male; 
from  whom  it  is  probable  that  these  have  been  transferred.  Lan- 
dois also  found  such  rudiments  on  the  under  surface  of  the  wing- 
covers  of  the  female  Achetidse,  and  on  the  femora  of  the  female 
Acridiidse.  In  the  Homoptera,  also,  the  females  have  the  proper 
musical  apparatus  in  a  functionless  state;  and  we  shall  hereafter 
meet  in  other  divisions  of  the  animal  kingdom  with  many  in- 
stances of  structures  proper  to  the  male  being  present  in  a  rudi- 
mentary condition  in  the  female. 


286  THE  DESCENT  OF  MAN. 

Landois  has  observed  another  important  fact,  namely,  that  in 
the  females  of  the  Acridiidae,  the  stridulating  teeth  on  the  femora 
remain  throughout  life  in  the  same  condition  in  which  they  first 
appear  during  the  larval  state  in  both  sexes.  In  the  males,  on 
the  other  hand,  they  become  further  developed,  and  acquire  their 
perfect  structure  at  the  last  moult,  when  the  insect  is  mature  and 
ready  to  breed. 

From  the  facts  now  given,  we  see  that  the  means  by  which 
the  males  of  the  Orthoptera  produce  their  sounds  are  extremely 
diversified,  and  are  altogether  different  from  those  employed  by 
the  Homoptera.*^  But  throughout  the  animal  kingdom  we  often 
find  the  same  object  gained  by  the  most  diversified  means;  this 
seems  due  to  the  whole  organization  having  undergone  multifari- 
ous changes  in  the  course  of  ages,  and  as  part  after  part  varied 
different  variations  were  taken  advantage  of  for  the  same  general 
purpose.  The  diversity  of  means  for  producing  sound  in  the  three 
families  of  the  Orthoptera  and  in  the  Homoptera,  impresses  the 
mind  with  the  high  importance  of  these  structures  to  the  males, 
for  the  sake  of  calling  or  alluring  the  females.  We  need  feel 
no  surprise  at  the  amount  of  modification  which  the  Orthoptera 
have  undergone  in  this  respect,  as  we  now  know,  from  Dr.  Scud- 
der's  remarkable  discovery,**  that  there  has  been  more  than  ample 
time.  This  naturalist  has  lately  found  a  fossil  insect  in  the 
Devonian  formation  of  New  Brunswick,  which  is  furnished  with 
"the  well-known  tympanum  or  stridulating  apparatus  of  the  male 
"Locustidae."  The  insect,  though  in  most  respects  related  to  the 
Neuroptera,  appears,  as  is  so  often  the  case  with  very  ancient 
forms,  to  connect  the  two  related  Orders  of  the  Neuroptera  and  Or- 
thoptera. 

I  have  but  little  more  to  say  on  the  Orthoptera.  Some  of  the 
species  are  very  pugnacious:  when  two  male  field-crickets 
(Gryllus  campestris)  are  confined  together,  they  fight  till  one  kills 
the  other;  and  the  species  of  Mantis  are  described  as  maneuvering 
with  their  sword-like  front-limbs,  like  hussars  with  their  sabres. 
The  Chinese  keep  these  insects  in  little  bamboo  cages,  and  match 
them  like  game-cocks.*^  With  respect  to  color,  some  exotic  locusts 
are  beautifully  ornamented;  the  posterior  wings  being  marked 
with  red,  blue,  and  black;  but  as  throughout  the  Order  the  sexes 
rarely  differ  much  in  color,  it  is  not  probable  that  they  owe  their 

*3  Landois  has  recently  found  in  certain  Orthoptera  rudimentary 
structures  closely  similar  to  the  sound-producing  organs  in  the  Ho- 
moptera; and  this  is  a  surprising  fact.  See  'Zeitschr,  fur  wissensch. 
Zoolog.'  B.  xxii.  Heft  3,  1871,  p.  348. 

**  'Transact.  Ent.  Soc'  3rd  series,  vol.  ii.  ('Journal  of  Proceedings," 
p.  117.) 

^  Westwood.  'Modem  Class,  of  Insects,'  vol.  i.  p.  427;  for  crickets, 
p.  445. 


ORTHOPTERA.  287 

bright  tints  to  sexual  selection.  Conspicuous  colors  may  be  of 
use  to  these  insects,  by  giving  notice  that  they  are  unpalatable. 
Thus  it  has  been  observed*^  that  a  bright-colored  Indian  locust 
was  invariably  rejected  when  offered  to  birds  and  lizards.  Some 
cases,  however,  are  known  of  sexual  differences  in  color  in  this 
Order.  The  male  of  an  American  cricket*^  is  described  as  being  as 
white  as  ivory,  whilst  the  female  varies  from  almost  white  to 
greenish-yellow  or  dusky.  Mr.  Walsh  informs  me  that  the  adult 
male  of  Spectrum  femoratum  (one  of  the  Phasmidse)  "is  of  a  shin- 
ning brownish-yellow  color;  the  adult  female  being  of  a  dull, 
"opaque,  cinereous  brown;  the  young  of  both  sexes  being  green." 
Lastly,  I  may  mention  that  the  male  of  one  curious  kind  of 
cricket*®  is  furnished  with  "a  long  membranous  appendage,  which 
"falls  over  the  face  like  a  veil;"  but  what  its  use  may  be,  is  not 
known. 

Order,  Neuroptera. — Little  need  here  be  said,  except  as  to  color. 
In  the  Ephemeridae  the  sexes  often  differ  slightly  in  their  ob- 
scure tints;*''  but  it  is  not  probable  that  the  males  are  thus  ren- 
dered attractive  to  the  females.  The  Libellulidse,  or  dragon-flies, 
are  ornamented  with  splendid  green,  blue,  yellow,  and  vermilion 
metallic  tints;  and  the  sexes  often  differ.  Thus,  as  Prof.  West- 
wood  remarks,^"  the  males  of  some  of  the  Agrionidas,  "are  of  a 
-■'rich  blue  with  black  wings,  whilst  the  females  are  fine  green  with 
"colorless  wings."  But  in  Agrion  Ramburii  these  colors  are  ex- 
actly reversed  in  the  two  sexes.^^  In  the  extensive  N.  American 
genus  of  Het^rina,  the  males  alone  have  a  beautiful  carmine  spot 
at  the  base  of  each  wing.  In  Anax  Junius  the  basal  part  of  the 
abdomen  in  the  male  is  a  vivid  ultramarine  blue  and  in  the  fe- 
male grass-green.  In  the  allied  genus  Gomphus,  on  the  other 
hand,  and  in  some  other  genera,  the  sexes  differ  but  little  in 
color.  In  closely-allied  forms  throughout  the  animal  king- 
dom, similar  cases  of  the  sexes  differing  greatly,  or  very 
little,  or  not  at  all,  are  of  frequent  occurrence.  Although  there  is 
so  wide  a  difference  in  color  between  the  sexes  of  many  Libel- 
lulidse, it  is  often  difficult  to  say  which  is  the  more  brilliant;  and 
the  ordinary  coloration  of  the  two  sexes  is  reversed,  as  we  have 
just  seen,  in  one  species  of  Agrion.    It  is  not  probable  that  their 

^8  Mr.  Ch.  Home,  in  'Proc.  Ent.  Soc'  May  3,  1869,  p.  xii. 

*'  The  Oecanthus  nivalis.  Harris,  'Insects  of  New  England,'  1842, 
p.  124.  The  two  sexes  of  Oe.  pellucidus  of  Europe  differ,  as  I  hear 
from  Victor  Carus,  in  nearly  the  same  manner. 

*8  Platyblemnus :     "Westwood,    'Modem  Class.'   vol.    i.    p.   447. 

*»  B.  D.  Walsh,  the  'Pseudo-neuroptera  of  Illinois,'  in  'Proc.  Ent. 
Soc.  of  Philadelphia,'  1862,  p.  361. 

so  'Modern  Class.'  vol.  ii.  p.  37. 

SI  Walsh,  ibid.  p.  381.  I  am  indebted  to  this  naturalist  for  the  fol- 
lowing facts  on  Hetaerina,  Anax,  and  Gomphus. 


2S8  THE  DESCENT  OF  MAN. 

colors  in  any  case  have  been  gained  as  a  protection.  Mr.  Mac- 
Lachlan,  who  has  closely  attended  to  this  family,  writes  to  me 
that  dragon-flies— tho  tyrants  of  the  Insect-world— are  the  least 
liable  of  any  insect  to  be  attacked  by  birds  or  other  enemies,  and 
he  believes  that  their  bright  colors  serve  as  a  sexual  attraction. 
Certain  dragon-flies  apparently  are  attracted  by  particular  colors: 
Mr.  Patterson  observed"  that  ihe  Agrionidae,  of  which  the  males 
are  blue,  settled  in  numbers  on  the  blue  float  of  a  fishing  line; 
whilst  two  other  species  were  attracted  by  shining  white  colors. 

It  is  an  interesting  fact,  first  noticed  by  Schelver,  that  in  sev- 
eral genera  belonging  to  two  sub-families,  the  males  on  first 
emergence  from  the  pupal  state,  are  colored  exactly  like  the 
females;  but  that  their  bodies  in  a  short  time  assume  a  con- 
spicuous milky-blue  tint,  owing  to  the  exudation  of  a  kind  of  oil, 
soluble  in  ether  and  alcohol.  Mr.  MacLachlan  believes  that  in 
the  male  of  Libellula  depressa  this  change  of  color  does  not  occur 
until  nearly  a  fortnight  after  the  metamorphosis,  when  the  sexes 
are  ready  to  pair. 

Certain  species  of  Neurothemis  present,  according  to  Brauer,''* 
a  curious  case  of  dimorphism,  some  of  the  females  having  ordinary 
wings,  whilst  others  have  them  "very  richly  netted,  as  in  the 
"males  of  the  same  species."  Brauer  "explains  the  phenomenon 
**on  Darwinian  principles  by  the  supposition  that  the  close  net- 
"ting  of  the  veins  is  a  secondary  sexual  character  in  the  males, 
"which  has  been  abruptly  transferred  to  some  of  the  females, 
"instead  of,  as  generally  occurs,  to  all  of  them."  Mr.  MacLachlan 
informs  me  of  another  instance  of  dimorphism  in  several  species 
of  Agrion,  in  which  some  individuals  are  of  an  orange  color,  and 
these  are  invariably  females.  This  is  probably  a  case  of  rever- 
sion; for  in  the  true  Libellulse,  when  the  sexes  differ  in  color, 
the  females  are  orange  or  yellow;  so  that  supposing  Agrion  to  be 
descended  from'some  primordial  form  which  resembled  the  typi- 
cal Libellulae  in  its  sexual  characters,  it  would  not  be  surprising 
that  a  tendency  to  vary  in  this  manner  should  occur  in  the  fe- 
males alone. 

Although  many  dragon-flies  are  large,  powerful,  and  fierce  in- 
sects, the  males  have  not  been  observed  by  Mr.  MacLachlan  to 
fight  together,  excepting,  as  he  believes,  in  some  of  the  smaller 
species  of  Agrion.  In  another  group  in  this  Order,  namely,  the 
Termites  or  white  ants,  both  sexes  at  the  time  of  swarming  may 
be  seen  running  about,  "the  male  after  the  female,  sometimes 
"two  chasing  one  female,  and  contending  with  great  eagerness 
"who  shall  win  the  prize."^*    The  Atropos  pulsatorius  is  said  to 

52  'Transact.  Ent.  Soc'  vol,   i.   1836,   p.  Ixxxi. 

53  See  abstract  in  the  'Zoological  Record,'  for  1867,  p.  450. 
"Kirby  and  Spence,  'Introduct.  to  Entomology,'  vol.  ii.  1818,  p.,  35. 


HYMENOPTERA.  289 

make  a  noise  with  its  jaws,  which  is  answered  by  other  indi- 
viduals/' 

Order,  Hymenoptera. — That  inimitable  observer,  M.  Fabre/«  in 
describing  the  habits  of  Cereeris,  a  wasp-like  insect,  remarks  that 
"fights  frequently  ensue  between  the  males  for  the  possession  of 
"some  particular  female,  who  sits  an  apparently  unconcerned  be- 
"holder  of  the  struggle  for  supremacy,  and  when  the  victory  is 
"decided,  quietly  flies  away  in  company  with  the  conqueror." 
Westwood^^  says  that  the  males  of  one  of  the  saw-flies  (Tenthre- 
din^)  "have  been  found  fighting  together  with  their  mandibles 
"locked."  As  M.  Fabre  speaks  of  the  males  of  Cereeris  striving 
to  obtain  a  particular  female,  it  may  be  well  to  bear  in  mind 
that  insects  belonging  to  this  Order  have  the  power  of  recognizing 
each  other  after  long  intervals  of  time,  and  are  deeply  attached. 
For  instance,  Pierre  Huber,  whose  accuracy  no  one  doubts,  sep- 
arated some  ants,  and  when,  after  an  interval  of  four  months, 
they  met  others  which  had  formerly  belonged  to  the  same  com- 
munity, they  recognized  and  caressed  one  another  with  their 
antennas.  Had  they  been  strangers  they  would  have  fought  to- 
gether. Again,  when  two  communities  engage  in  a  battle,  the 
ants  on  the  same  side  sometimes  attack  each  other  in  the  general 
confusion,  but  they  soon  perceive  their  mistake,  and  the  one  ant 
soothes  the  other.^® 

In  this  Order  slight  differences  in  color,  according  to  sex,  are 
common,  but  conspicuous  differences  are  rare  except  in  the  fam- 
ily of  Bees;  yet  both  sexes  of  certain  groups  are  so  brilliantly 
colored — for  instance  in  Chrysis,  in  which  vermilion  and  metallic 
greens  prevail — that  we  are  tempted  to  attribute  the  result  to 
sexual  selection.  In  the  Ichneumonidse,  according  to  Mr.  Walsh,^' 
the  males  are  almost  universally  lighter-colored  than  the  females. 
On  the  other  hand,  in  the  Tenthredinidae  the  males  are  generally 
darker  than  the  females.  In  the  Siricidae  the  sexes  frequently  dif- 
fer; thus  the  male  of  Sirex  juvencus  is  banded  with  orange, 
whilst  the  female  is  dark  purple;  but  it  is  diflacult  to  say  which 
sex  is  the  more  ornamented.  In  Tremex  columbae  the  female  is 
much  brighter  colored  than  the  male.  I  am  informed  by  Mr.  F. 
Smith,  that  the  male  ants  of  several  species  are  black,  the  females 
being  testaceous. 

In  the  family  of  Bees,  especially  in  the  solitary  species,  as  I 

^  Houzeau,   'Les  Facultes  Men  tales,'  &c.     Tom.  i.  p.  104. 

^  See  an  interesting  article,  'The  Writings  of  Fabre,'  in  'Nat.  Hist. 
Review,'  April,  1862,  p.  122. 

E7  'Journal  of  Proc.  of  Entomolog.  Soc'  Sept.  7th,  1863,  p.  169. 

^  P.  Huber,  'Recherches  sur  les  Moeurs  des  Fourmis,'  1810,  pp.  ISflt 
165. 

5»  'Proc.  Entomolog-.  Soc.  of  Philadelphia,'  1866,  pp.  238-239. 
20 


290  THE  DESCENT  OF  MAN. 

hear  from  the  same  entomologist,  the  sexes  often  differ  in  color. 
The  males  are  generally  the  brighter,  and  in  Bombus  as  well  as  in 
Apathus,  much  more  variable  in  color  than  the  females.  In 
Anthophora  retusa  the  male  is  of  a  rich  fulvous-brown,  whilst  the 
female  is  quite  black;  so  are  the  females  of  several  species  of 
Xylocopa,  the  males  being  bright  yellow.  On  the  other  hand  the 
females  of  some  species,  as  of  Andrsena  fulva,  are  much  brighter- 
colored  than  the  males.  Such  differences  in  color  can  hardly  be 
accounted  for  by  the  males  being  defenseless  and  thus  requiring 
protection,  whilst  the  females  are  well  defended  by  their  stings. 
H.  Muller,"^"  who  has  particularly  attended  to  the  habits  of  bees, 
attributes  these  differences  in  color  in  chief  part  to  sexual  selec- 
tion. That  bees  have  a  keen  perception  of  color  is  certain.  He 
says  that  the  males  search  eagerly  and  fight  for  the  possession 
of  the  females;  and  he  accounts  through  such  contests  for  the 
mandibles  of  the  males  being  in  certain  species  larger  than  those 
of  the  females.  In  some  cases  the  males  are  far  more  numerous 
than  the  females,  either  early  in  the  season,  or  at  all  times  and 
places,  or  locally;  whereas  the  females  in  other  cases  are  appar- 
ently in  excess.  In  some  species  the  more  beautiful  males  appear 
to  have  been  selected  by  the  females;  and  in  others  the  more  beau- 
tiful females  by  the  males.  Consequently  in  certain  genera  (Miil- 
ler,  p.  42),  the  males  of  the  several  species  differ  much  in  appear- 
ance, whilst  the  females  are  almost  indistinguishable;  in  other 
genera,  the  reverse  occurs.  H.  Miiller  believes  (p.  82)  that  the 
colors  gained  by  one  sex  through  sexual  selection  have  often  been 
transferred  in  a  variable  degree  to  the  other  sex,  just  as  the  pollen- 
collecting  apparatus  of  the  female  has  often  been  transferred  to 
the  male,  to  whom  it  is  absolutely  useless.^^ 


6»  'Anwendung  der  Darwinschen  Lehre  auf  Bienen.'  Verb.  d.  n. 
Jahrg.  xxix. 

61 M.  Perrier  in  his  article  'la  Selection  sexuelle  d'apres  Darwin* 
('Revue  Scientifique,'  Feb.  1873,  p.  868),  without  apparently  having-  re- 
flected much  on  the  subject,  objects  that  as  the  males  of  social  bees 
are  known  to  be  produced  from  unfertilized  ova,  they  could  not  trans- 
mit new  characters  to  their  male  offspring.  This  is  an  extraordinary 
objection.  A  female  bee  fertilized  by  a  male,  which  presented  some 
character  facilitating  the  union  of  the  sexes,  or  rendering  him  more 
attractive  to  the  female,  would  lay  eggs  which  would  produce  only 
females;  but  these  young  females  would  next  year  produce  males; 
and  will  it  be  pretended  that  such  males  would  not  inherit  the  char- 
acters of  their  male  grandfathers?  To  take  a  case  with  ordinary 
animals  as  nearly  parallel  as  possible:  if  a  female  of  any  white 
quadruped  or  bird  were  crossed  by  a  male  of  a  black  breed,  and  the 
male  and  female  offspring  were  paired  together,  will  it  be  pretended 
that  the  grandchildren  would  not  inherit  a  tendency  to  blackness 
from  their  male  grandfather?  The  acquirement  of  new  characters 
by  the  sterile  warker-bees  is  a  much  more  difficult  case,  but  I  have 


COLEOPTERA.  291 

Mutilla  EuropaBa  makes  a  stridulating  noise;  and  according  to 
Goureau^^  both  sexes  have  this  power.  He  attributes  the  sound 
to  the  friction  of  the  third  and  preceding  abdominal  segments, 
and  I  find  that  these  surfaces  are  marked  with  very  fine  con- 
centric ridges;  but  so  is  the  projecting  thoracic  collar,  into  which 
the  head  articulates,  and  this  collar,  when  scratched  with  the 
point  of  a  needle,  emits  the  proper  sound.  It  is  rather  surprising 
that  both  sexes  should  have  the  power  of  stridulating,  as  the 
male  is  winged  and  the  female  wingless.  It  is  notorious  that 
Bees  express  certain  emotions,  as  of  anger,  by  the  tone  of  their 
humming;  and  according  to  H.  Miiller  (p.  80),  the  males  of  some 
species  makes  a  peculiar  singing  noise  wnilst  pursuing  the  females. 

Order,  Coleoptera  (Beetles). — Many  beetles  are  colored  so  as 
to  resemble  the  surfaces  which  they  habitually  frequent,  and 
they  thus  escape  detection  by  their  enemies.  Other  species,  for 
instance  diamond-beetles,  are  ornamented  with  splendid  colors, 
which  are  often  arranged  in  stripes,  spots,  crosses,  and  other 
elegant  patterns.  Such  colors  can  hardly  serve  directly  as  a 
protection,  except  in  the  case  of  certain  flower-feeding  species; 
but  they  may  serve  as  a  warning  or  means  of  recognition,  on  the 
same  principle  as  the  phosphorescence  of  the  glow-worm.  As 
with  beetles  the  colors  of  the  two  sexes  are  generally  alike,  we 
have  no  evidence  that  they  have  been  gained  through  sexual 
selection;  but  this  is  at  least  posible,  for  they  may  have  been 
developed  in  one  sex  and  then  transferred  to  the  other;  and 
this  view  is  even  in  some  degree  probable  in  those  groups  which 
possess  other  well-marked  secondary  sexual  characters.  Blind 
beetles,  which  cannot  of  course  behold  each  other's  beauty,  never, 
as  I  hear  from  Mr.  Waterhouse,  jun.,  exhibit  bright  colors, 
though  they  often  have  polished  coats;  but  the  explanation  of 
their  obscurity  may  be  that  they  generally  inhabit  caves  and 
other  obscure  stations. 

Some  Longicorns,  especially  certain  Prionid^,  offer  an  excep- 
tion to  the  rule  that  the  sexes  of  beetles  do  not  differ  in  color. 
Most  of  these  insects  are  large  and  splendidly  colored.  The 
males  in  the  genus  Pyrodes,**^  which  I  saw  in  Mr.  Bates's  collec- 

endeavored  to  show  in  my  'Origin  of  Species,'  how  these  sterile  be- 
ings are  subjected  to  the  power  of  natural  selection. 

^2  Quoted  by  Westwood,  'Modern  Class,  of  Insects,'  vol.  ii.  p.  214. 

^  Pyrodes  pulcherrimus,  in  which  the  sexes  differ  conspicuously, 
has  been  described  by  Mr.  Bates  in  'Transact.  Ent.  Soc'  1869,  p.  50.  I 
will  specify  the  few  other  cases  in  which  I  have  heard  of  a  differ- 
ence in  color  between  the  sexes  of  beetles.  Kirby  and  Spence  ('In- 
troduct.  to  Entomology,'  vol.  iii.  p.  301)  mention  a  Cantharis,  Meloe, 
Ehagium,  and  the  Leptura  testacea;  the  male  of  the  latter  being  tes- 
taceous, with  a  black  thorax,  and  the  female  of  a  dull  red  all  over. 
These  two  latter  beetles  belong  to  the  family  of  Longicorns.     Messrs. 


292 


THE  DESCENT  OF  MAN. 


tion,  are  generally  redder  but  rather  duller  than  the  females,  the 
latter  being  colored  of  a  more  or  less  splendid  golden-green.  On 
the  other  hand,  in  one  species  the  male  is  golden-green,  the  female 
being  richly  tinted  with  red  and  purple.  In  the  genus  Esmeralda 
the  sexes  differ  so  greatly  in  color  that  they  have  been  ranked 
as  distinct  species;  in  one  species  both  are  of  a  beautiful  shin- 
ing green,  but  the  male  has  a  red  thorax.    On  the  whole,  as  far 


Fig.  16.    Chalcosoma    atlas.      Upper    figure,    male    (reduced);    lower 
figure,   female   (natural   size). 

as  I  could  judge,  the  females  of  those  Prionidse,  in  which  the  sexes 
differ,  are  colored  more  richly  than  the  males,  and  this  does  not 
accord  with  the  common  rule  in  regard  to  color,  when  acquired 
through  sexual  selection. 

A  most  remarkable  distinction  between  the  sexes  of  many 
beetles  is  presented  by  the  great  horns  which  rise  from  the  head, 
thorax,  and  clypeus  of  the  males;  and  in  some  few  cases  from 
the  under  surface  of  the  body.  These  horns,  in  the  great  family 
of  the  Lamellicorns,  resemble  these  of  various  quadrupeds,  such 


R.  Trim  en  and  Waterhouse,  jun.,  inform  me  of  two  Lamellicorns, 
viz.,  a  Peritrichia  and  Triohius,  the  male  of  the  latter  being  more 
obscurely  colored  than  the  female.  In  Tillus  elongatus  the  male  is 
black,  and  the  female  always,  as  it  is  believed,  of  a  dark  blue  color, 
with  a  red  thorax.  The  male,  also,  of  Orsodacna  atra,  as  I  hear  from 
Mr.  Walsh,  is  black,  the  female  (the  so-called  O.  ruficoUis)  having  a 
rufous  thorax. 


COLEOPTERA. 


293 


Fig-.  17.  Copris    isidis.      (Left-hand  figures,  males.) 


Fig.  20. 


Onthophagus    rangifer,    enlarged. 


'K) 


294  TK23  DESCENT  OF  MAN. 

as  stags,  rhinoceroses,  &c.,  and  are  wonderful  both  from  their 
size  and  diversified  shapes.  Instead  of  describing  them,  I  have 
given  figures  of  the  males  and  females  of  some  of  the  more  re- 
markable forms.  (Figs.  16  to  20.)  The  females  generally  ex- 
hibit rudiments  of  the  horns  in  the  form  of  small  knobs  or  ridges; 
but  some  are  destitute  of  even  the  slightest  rudiment.  On  the 
other  hand,  the  horns  are  nearly  as  well  developed  in  the  female 
as  in  the  male  of  Phanasus  lancifer;  and  only  a  little  less  well 
developed  in  the  females  of  some  other  species  of  this  genus  and 
of  Copris.  I  am  informed  by  Mr.  Bates  that  the  horns  do  not 
differ  in  any  manner  corresponding  with  the  more  important  char- 
acteristic differences  between  the  several  subdivisions  of  the 
family:  thus  within  the  same  section  of  the  genus  Onthophagus, 
there  are  species  which  have  a  single  horn,  and  others  which 
have  two. 

In  almost  all  cases,  the  horns  are  remarkable  from  their  ex- 
cessive variability;  so  that  a  graduated  series  can  be  formed, 
from  the  most  highly  developed  males  to  others  so  degenerate 
that  they  can  barely  be  distinguished  from  the  females.  Mr. 
Walsh^*  found  that  in  Phanaeus  carnifex  the  horns  were  thrice  as 
long  in  some  males  as  in  others.  Mr.  Bates,  after  examining 
above  a  hundred  males  of  Onthophagus  rangifer  (fig.  20),  thought 
that  he  had  at  last  discovered  a  species  in  which  the  horns  did 
not  vary;    but  further  research  proved  the  contrary. 

The  extraordinary  size  of  the  horns,  and  their  widely  different 
structure  in  closely-allied  forms,  indicate  that  they  have  been 
formed  for  some  purpose;  but  their  excessive  variability  in  the 
males  of  the  same  species  leads  to  the  inference  that  this  purpose 
cannot  be  of  a  definite  nature.  The  horns  do  not  show  marks  of 
friction,  as  if  used  for  any  ordinary  work.  Some  authors  sup- 
pose®^ that  as  the  males  wander  about  much  more  than  the  fe- 
males, they  require  horns  as  a  defense  against  their  enemies;  but 
as  the  horns  are  often  blunt,  they  do  not  seem  well  adapted  for 
defense.  The  most  obvious  conjecture  is  that  they  are  used  by 
the  males  for  fighting  together;  but  the  males  have  never  been 
observed  to  fight;  nor  could  Mr.  Bates,  after  a  careful  exam- 
ination of  numerous  species,  find  any  suflicient  evidence,  in  thei^ 
mutilated  or  broken  condition,  of  their  having  been  thus  used. 
If  the  males  had  been  habitual  fighters,  the  size  of  their  bodies 
would  probably  have  been  increased  through  sexual  selection, 
so  as  to  have  exceeded  that  of  the  females;  but  Mr.  Bates,  after 
comparing  the  two  sexes  in  above  a  hundred  species  of  the 
Copridae,  did  not  find  any  marked  difference  in  this  respect 
amongst  well-developed  individuals.    In  Lethrus,  moreover,  a  bee- 

«*  'Proc.  Entomolog.   Soc.  of  Philadelphia,'  1864,  p.  228. 

65  Kirby  and  f-ipence,  'Introduct.  Entomolog-.'  vol.  iii.  p.  300. 


COLEOPTERA. 


295 


tie  belonging  to  the  same  great  division  of  the  Lamellicorns,  the 
males  are  known  to  fight,  but  are  not  provided  with  horns,  though 
their  mandibles  are  much  larger  than  those  of  the  female. 

The  conclusion  that  the  horns  have  been  acquired  as  ornaments 
is  that  which  best  agrees  with  the  fact  of  their  having  been  so 
immensely,  yet  not  fixedly,  developed,— as  shown  by  their  extreme 
variability  in  the  same  species,  and  by  their  extreme  diversity  in 
closely-allied  species.  This  view  will  at  first  appear  extremely 
improbable;  but  we  shall  hereafter  find  with  many  animals  stand- 
ing much  higher  in  the  scale,  namely  fishes,  amphibians,  reptiles 
and  birds,  that  various  kinds  of  crests,  knobs,  horns  and  combs 
have  been  developed  apparently  for  this  sole  purpose. 

The  males  of  Onitis  furcifer  (fig.  21),  and  of 
some  other  .species  of  the  genus,  are  furnished 
with  singular  projections  on  their  anterior  fe- 
mora, and  with  a  great  fork  or  pair  of  horns  on 
the  lower  surface  of  the  thorax.  Judging  from 
other  insects,  these  may  aid  the  male  in  clinging 
to  the  female.  Although  the  males  have  not 
even  a  trace  of  a  horn  on  the  upper  surface  of 
the  body,  yet  the  females  plainly  exhibit  a  rudi- 
21.  Onitis  ^^^^  of  a  single  horn  on  the  head  (fig.  22,  a), 
male  and  of  a  crest  (b)  on  the  thorax.  That  the 
slight  thoracic  crest  in  the  female  is  a  rudiment 
of  a  projection  proper  to  the  male,  though  en- 
tirely absent  in  the  males  of  this  particular  species,  is  clear: 
for  the  female  of  Bubas  bison  (a  genus  which  comes 
next    to    Onitis)    has    a    similar    slight    crest    on    the    thorax. 


Fig. 
furcifer, 
viewed  from  be 
neath. 


Fig.  22.    Left-hand  figure,  male  of  Onitis  furcifer,  viewed  laterally. 

Right-hand  figure,   female,     a.  Rudiment  of   cephalic  horn. 

b.  Trace  of  thoracic  horn  or  crest. 

and  the  male  bears  a  great  projection  in  the  same  situa- 
tion. So,  again,  there  can  hardly  be  a  doubt  that  the  little  point 
(a)  on  the  head  of  the  female  Onitis  furcifer,  as  well  as  on  the 
head  of  the  females  of  two  or  three  allied  species,  is  a  rudimentary 
representative  of  the  cephalic  horn,  which  is  common  to  the 
males  of  so  many  Lamellicorn  beetles,  as  in  Ph^tnseus  (fig.  18). 
The  old  belief  that  rudiments  have  been  created  to  complete  the 


296  THE  DESCENT  OF  MAN. 

scheme  of  nature  is  here  r^o  far  from  holding  good,  that  we  have 
a  complete  inversion  of  the  ordinary  state  of  things  in  the  family. 
We  may  reasonably  suspect  that  the  males  originally  bore  horns 
and  transferred  them  to  the  females  in  a  rudimentary  condition, 
as  in  so  many  other  Lamellicorns.  Why  the  males  subsequently 
lost  their  horns,  we  know  not;  but  this  may  have  been  caused 
through  the  principle  of  compensation,  owing  to  the  development 
of  the  large  horns  and  projections  on  the  lower  surface;  and  as 
these  are  confined  to  the  males,  the  rudiments  of  the  upper  horns 
on  the  females  would  not  have  been  thus  obliterated. 

The  cases  hitherto  given  refer  to  the  Lamellicorns,  but  the  males 
of  some  few  other  beetles,  belonging  to  two  widely  distinct  groups, 
namely,  the  Curculionidse  and  Staphylinidae,  are  furnished  with 
horns — in  the  former  on  the  lower  surface  of  the  body,*^^  in  the 
latter  on  the  upper  surface  of  the  head  and  thorax.  In  the  Staphy- 
linidse,  the  horns  of  the  males  are  extraordinarily  variable  in  the 
same  species,  just  as  we  have  seen  with  the  Lamellicorns.  In 
Siagonium  we  have  a  case  of  dimorphism,  for  the  males  can  be 
divided  into  two  sets,  differing  greatly  in  the  size  of  their  bodfes 
and  in  the  development  of  their  horns,  without  intermediate 
gradations.    In  a  species  of  Bledius  (fig.  23),  also  belonging  to  the 


Fig.   23.     Bledius   taurus,    magnified.     Left-hand   figure,   male;    right- 
hand  figure,  female. 

Staphylinidae,  Professor  Westwood  states  that,  "male  specimens 
"can  be  found  in  the  same  locality  in  which  the  central  horn  of 
"the  thorax  is  very  large,  but  the  horns  of  the  head  quite  rudi- 
"mental;  and  others,  in  which  the  thoracic  horn  is  much  shorter, 
"whilst  the  protuberances  on  the  head  are  long.""  Here  we  ap- 
parently have  a  case  of  compensation,  which  throws  light  on  that 
just  given  of  the  supposed  loss  of  the  upper  horns  by  the  males  of 
Onitis. 

Law  of  Battle. -^Some  male  beetles,  which  seem  ill-fitted  for 
fighting,  nevertheless  engage  in  conflicts  for  the  possession  of  the 
females.    Mr.  Wallace®*  saw  two  males  of  Leptorhynchus  angusta- 

«« Kirby  and  Spence,  'Introduct.  Entomolog.'  vol.  iii.  p.  329. 

^7  'Modern  Classification  of  Insects,'  vol.  i.  p.  172:  Siagonium,  p.  172. 
In  the  British  Museum  I  noticed  one  male  specimen  of  Siagonium  in 
an  intermediate  condition,  so  that  the  dimorphism  is  not  strict. 

«8  'The  Malay  Archipelago,'  vol.  ii.  1869,  p.  276.  Riley,  Sixth  'Re- 
port on  insects  of  Missouri,'  1874,  p.  115. 


COLEOPTERA.  297 

tus,  a  linear  beetle  with  a  much  elongated  rostrum,  "fighting  for 
"a  female,  who  stood  close  by  busy  at  her  boring.  They  pushed 
"at  each  other  with  their  rostra,  and  clawed  and  thumped,  ap- 
"parently  in  the  greatest  rage."  The  smaller  male,  however, 
"soon  ran  away,  acknowledging  himself  vanquished."  In  some 
few  cases  male  beetles  are  well  adapted  for  fighting,  by  possess- 
ing great  toothed  mandibles,  much  larger  than  those  of  the  fe- 
males. This  is  the  case  with  the  common  stag-beetle  (Lucanus 
cervus),  the  males  of  which  emerge  from  the  pupal  state  about  a 
week  before  the  other  sex,  so  that  several  may  often  be  seen 
pursuing  the  same  female.  At  this  season  they  engage  in  fierce 
conflicts.  When  Mr.  A.  H.  Davis*^*  enclosed  two  males  with  one 
female  in  a  box,  the  larger  male  severely  pinched  the  smaller  one, 
until  he  resigned  his  pretensions.  A  friend  informs  me  that  when 
a  boy  he  often  put  the  males  together  to  see  them  fight,  and  he 
noticed  that  they  were  much  bolder  and  fiercer  than  the  females, 
as  with  the  higher  animals.  The  males  would  seize  hold  of  his 
finger,  if  held  in  front  of  them,  but  not  so  the  females,  although 
they  have  stronger  jaws.  The  males  of  many  of  the  Lucanidse,  as 
well  as  of  the  above-mentioned  Leptorhynchus,  are  larger  and 
more  powerful  insects  than  the  females.  The  two  sexes  of  Le- 
thrus  cephalotes  (one  of  the  Lamellicorns)  inhabit  the  same  bur- 
row; and  the  male  has  larger  mandibles  than  the  female.  If,  dur- 
ing the  breeding-season,  a  strange  male  attempts  to  enter  the  bur- 
row, he  is  attacked;  the  female  does  not  remain  passive,  but  closes 
the  mouth  of  the  burrow,  and  encourages  her  mate  by  continually 
pushing  him  on  from  behind;  and  the  battle  lasts  until  the  ag- 
gressor is  killed  or  runs  away.^°  The  two  sexes  of  another 
Lamellicorn  beetle,  the  Ateuchus  cicatricosus,  live  in  pairs,  and 
seem  much  attached  to  each  other;  the  male  excites  the  female  to 
roll  the  ball  of  dung  in  which  the  ova  are  deposited;  and  if  she  is 
removed,  he  becomes  much  agitated.  If  the  male  is  removed  the 
female  ceases  all  work,  and  as  M.  Brulerie'^  believes,  would  remain 
on  the  same  spot  until  she  died. 

The  great  mandibles  of  the  male  Luscanidse  are  extremely  va- 
riable both  in  size  and  structure,  and  in  this  respecet  resemble 
the  horns  on  the  head  and  thorax  of  many  male  Lamellicorns  and 
Staphylinidae.  A  perfect  series  can  be  formed  from  the  best-pro- 
vided to  the  worst-provided  or  degenerate  males.  Although  the 
mandibles  of  the  common  stag-beetle,  and  probably  of  many  other 
species,  are  used  as  efficient  weapons  for  fighting,  it  is  doubtful 

«» 'Entomological  Magazine,'  vol.  i.  1833,  p.  82.  See,  also,  on  the 
conflicts  of  this  species,  Kirby  and  Spence,  ibid,  vol.  iii.  p.  314;  and 
Westwood,  ibid.  vol.  i.  p.  187. 

^0  Quoted  from  Fischer,  in  'Diet.  Class.  d'Hist.  Nat.'  tom.  x.  p.  324. 

7^  'Ann.  Soc.  Entomolog.  France,'  1866,  as  quoted  in  'Journal  of 
Travel,'  by  A.  Murray,  1868,  p.  135. 


298 


THE  DESCENT  OF  MAN. 


whether  their  great  size  can  thus  be  accounted  for.  We  have  seen 
that  they  are  used  by  the  Lucanus  elaphus  of  N.  America  for  seiz- 
ing the  female.  As  they  are  so  conspic- 
uous and  so  elegantly  branched,  and  as 
owing  to  their  great  length  they  are  not 
well  adapted  for  pinching,  the  suspicion 
has  crossed  my  mind  that  they  may  in 
addition  serve  as  an  ornament,  like  the 
horns  on  the  head  and  thorax  of  the 
various  species  above  described.  The 
male  Chiasognathus  Grantii  of  S.  Chile 
— a  splendid  beetle  belonging  to  the 
same  family — has  enormously  developed 
mandibles  (iig.  24) ;  he  is  bold  and  pugna- 
cious; when  threatened  he  faces  round, 
opens  his  great  jaws,  and  at  the  same 
time  stridulates  loudly.  But  the  mandi- 
bles were  not  strong  enough  to  pinch  my 
linger  so  as  to  cause  actual  pain. 

Sexual  selection,  which  implies  the  pos- 
session of  considerable  perceptive  powers 
and  of  strong  passions,  seems  to  have 
been  more  effective  with  the  Lamellicorna 
than  with  any  other  family  of  beetles. 
With  bome  species  the  males  are  provided 
J     p  \  with  weapons  for  fighting;    some  live  in 

^   if         ^  I  pairs  and  show  mutual  affection;    many 

have  the  power  of  stridulating  when  ex- 
cited; many  are  furnished  with  the  most 
extraordinary  horns,  apparently  for  the 
sake  of  ornament;  and  some,  which  are 
diurnal  in  their  habits,  are  gorgeously 
colored.  Lastly,  several  of  the  largest 
beetles  in  the  world  belong  to  this  fam- 
ily, which  was  placed  by  Linnaeus  and 
Fabricius  at  the  head  of  the  Order.^^ 


Fig".    24.     Chiasognathus 


Stridulating  organs. — Beetles  belonging 
to  many  and  widely  distinct  families  pos- 
sess these  organs.     The  sound  thus  pro- 


grantii,  reduced, 
per  figrire,  male; 
er  figure,  female. 


Up-   duced  can  sometimes  be  heard  at  the  dis- 


low- 


tance    of   several    feet    or    even    yards,''^ 
but    it    is    not    comparable    with    that 


made    by    the    Orthoptera.       The    rasp    generally    consists    of 


'2  Westwood,  'Modern  Class.'  vol.  1.  p.  184. 

7s  Wollaston,   'On  certain  Musical  Curculionidae,'   'Annals  and  Mag. 
of  Nat.  Hist.,  vol.  vi.  1860,  p.  14. 


COLEOPTERA. 


2^9 


a  narrow,  slightly-raised  surface,  crossed  by  very  fine,  par- 
allel ribs,  sometimes  so  fine  as  to  cause  irridescent  colors, 
and  having  a  very  elegant  appearance  under  the  micro- 
scope. In  some  cases,  as  with  Typhceus,  minute,  bristly  or 
scale-like  prominences,  with  which  the  whole  surrounding  surface 
is  covered  in  approximately  parallel  lines,  could  be  traced  passing 
into  the  ribs  of  the  rasp.  The  transition  takes  place  by  their  be- 
coming confluent  and  straight,  and  at  the  same  time  more  promi- 
nent and  smooth.  A  hard  ridge  on  an  adjoining  part  of  the  body 
serves  as  the  scraper  for  the  rasp,  but  this  scraper  in  some  cases 
has  been  specially  modified  for  the  purpose.  It  is  rapidly  moved 
across  the  rasp,  or  conversely  the  rasp  across  the  scraper. 


Fiff.  25. 


Necrophorus  (from  Landois).    r.  The  two  rasps, 
figure,  part  of  the  rasp  highly  magnified. 


Left-hand 


These  organs  are  situated  in  widely  different  positions.  In  the 
carrion-beetles  (Necrophorus)  two  parallel  rasps  (r,  fig.  25)  stand 
on  the  dorsal  surface  of  the  fifth  abdominal  segment,  each  rasp'* 
consisting  of  126  to  140  fine  ribs.  These  ribs  are  scraped  against 
the  posterior  margins  of  the  elytra,  a  small  portion  of  which  pro- 
jects beyond  the  general  outline.  In  many  Crioceridae,  and  in 
Clythra  4-punctata  (one  of  the  Chrysomelidse),  and  in  some  Tene- 
brionidae,  &c.,''^  the  rasp  is  seated  on  the  dorsal  apex  of  the  abdo- 
men, on  the  pygidium  or  pro-pygidium,  and  is  scraped  in  the  same 
manner  by  the  elytra.  In  Heterocerus,  which  belongs  to  another 
family,  the  rasps  are  placed  on  the  sides  of  the  first  abdominal 


7*  Landois,  'Zeitschrift  fur  wiss.  Zoolog.'  B.  xvii.  1867,  s.  127. 

■^5  I  am  greatly  indebted  to  Mr.  G.  R.  Crotch  for  having  sent  me 
many  prepared  specimens  of  various  beetles  belonging  to  these  three 
families  and  to  others,  as  well  as  for  valuable  information.  He  be- 
lieves that  the  power  of  stridulation  in  the  Clythra  has  not  been  pre- 
viously observed.  I  am  also  much  indebted  to  Mr.  E.  W.  Janson, 
for  information  and  specimens.  I  may  add  that  my  son,  Mr.  P.  Dar- 
win, finds  that  Dermestes  murinus  stridulates,  but  he  searched  in 
vain  for  the  apparatus.  Scolytus  has  lately  been  described  by  Dr. 
Chapman  as  a  stridulator,  in  the  'Entomologist's  Monthly  Magazine,* 
vol.  vi.  p.  130. 


300 


THE  DESCENT  OF  MAN. 


segment,  and  are  scraped  by  ridges  on  the  femora.'*'  In  certain 
Curcullonidse  and  Carabidae,"  the  parts  are  completely  reversed  in 
position,  for  the  rasps  are  seated  on  the  inferior  surface  of  the 
elytra,  near  their  apices,  or  along  their  outer  margins,  and  the 
edges  of  the  abdominal  segments  serve  as  the  scrapers.  In  Pelo- 
bius  Hermann!  (one  of  Dytiscidse  or  water-beetles)  a  strong  ridge 
runs  parallel  and  near  to  the  sutural  margin  of  the  elytra,  and  is 
crossed  by  ribs,  coarse  in  the  middle  part,  but  becoming  gradually 
finer  at  both  ends,  especially  at  the  upper  end;  when  this  insect  is 
held  under  water  or  in  the  air  a  stridulating  noise  is  produced 
by  the  extreme  horny  margin  of  the  abdomen  being  scraped 
against  the  rasps.  In  a  great  number  of  long-horned  beetles 
(Longicornia)  the  organs  are  situated  quite  otherwise,  the  rasp 
being  on  the  meso-thorax,  which  is  rubbed  against  the  pro-thorax; 
Landois  counted  238  very  fine  ribs  on  the  rasp 
jf  of  Cerambyx  heros. 

Many  Lamellicorns  have  the  power  of  stridu- 
lating, and  the  organs  differ  greatly  in  position. 
Some  species  stridulate  very  loudly,  so  that 
when  Mr.  F.  Smith  caught  a  Trox  sabulosus, 
a  gamekeeper,  who  stood  by,  thought  he  had 
caught  a  mouse;  but  I  failed  to  discover  the 
proper  organs  in  this  beetle.  In  Geotrupes  and 
Typhosus  a  narrow  ridge  runs  obliquely  across 
(r.  fig.  26)  the  coxa  of  each  hind-leg  (having  in 
G.  stercorarius  84  ribs),  which  is  scraped  by  a 
specially  projecting  part  of  one  of  the  abdominal 
segments.  In  the  nearly  allied  Copris  lunaris, 
an  excessively  narrow  fine  rasp  runs  along  the 
sutural  margin  of  the  elytra,  with  another  short 
rasp  near  the  basal  outer  margin;  but  in  some 
other  Coprini  the  rasp  is  seated,  according  to 
Leconte,-^  on  the  dorsal  surface  of  the  abdomen. 
In  Oryctes  it  is  seated  on  the  pro-pygidium;  and, 
according  to  the  same  entomologist,  in  some 
other  Dynastini,  on  the  under  surface  of  the  elytra.    Lastly,  West- 


Fig.  26.  Hind- 
leg  of  Geo- 
trupes stercor- 
arius (from 
Landois). 

r.  Rasp.  c.  Coxa, 
f.  Femur,  t. 
Tibia  tr.  Tarsi. 


'6  Schiodte,  translated  in  'Annals  and  Mag.  of  Nat.  Hist.'  vol.  xx. 
1867,  p.  37. 

"  Westring  has  described  (Kroyer,  'Naturhist.  Tidskrift,'  B.  ii.  1848- 
49,  p.  334)  the  stridulating  organs  in  these  two,  as  well  as  in  other 
families.  In  the  Carabidae  I  have  examined  Elaphrus  uliginosus  and 
Blethisa  multipunctata,  sent  to  me  by  Mr.  Crotch.  In  Blethisa  the 
transverse  ridges  on  the  furrowed  border  of  the  abdominal  segment 
do  not,  as  far  as  I  could  judge,  come  into  play  in  scraping  the  rasps 
on  the  elytra. 

78  1  am  indebted  to  Mr,  Walsh,  of  Illinois,  for  having  sent  me  ex- 
tracts from  Leconte's  'Introduction  to  Entomology,'  pp.  101,  143. 


COLEOPTERA.  301 

ring  states  that  in  Omaloplia  brunnea  the  rasp  is  placed  on  the  pro- 
sternum,  and  the  scraper  on  the  meta-stei'num,  the  parts  thus  oc- 
cupying the  under  surface  of  the  body,  instead  of  the  upper  sur- 
face as  in  the  Longicorns. 

We  thus  see  that  in  the  different  coleopterous  families  the 
stridulating  organs  are  wonderfully  diversified  in  position,  but  not 
much  in  structure.  Within  the  same  family  some  species  are  pro- 
vided with  these  organs,  and  others  are  destitute  of  them.  This 
diversity  is  intelligible,  if  we  suppose  that  originally  various  bee- 
tles made  a  shuffling  or  hissing  noise  by  the  rubbing  together  of 
any  hard  and  rough  parts  of  their  bodies,  which  happened  to  be  in 
contact;  and  that  from  the  noise  thus  produced  being  in  some  way 
useful,  the  rough  surfaces  were  gradually  developed  into  regular 
stridulating  organs.  Some  beetles  as  they  move,  now  produce, 
either  intentionally  or  unintentionally,  a  shuffling  noise,  without 
possessing  any  proper  organs  for  the  purpose.  Mr.  Wallace  in- 
forms me  that  the  Euchirus  longimanus  (a  Lamellicorn,  with  the 
anterior  legs  wonderfully  elongated  in  the  male)  "makes,  whilst 
"moving,  a  low  hissing  sound  by  the  protrusion  and  contraction  of 
"the  abdomen;  and  when  seized  it  produces  a  grating  sound  by 
"rubbing  its  hind-legs  against  the  edges  of  the  elytra."  The  hiss- 
ing sound  is  clearly  due  to  a  narrow  rasp  running  along  the  sut- 
ural  margin  of  each  elytron;  and  I  could  likewise  make  the  grat- 
ing sound  by  rubbing  the  shagreened  surface  of  the  femur  against 
the  granulated  margin  of  the  corresponding  elytron;  but  I  could 
not  here  detect  any  proper  rasp;  nor  is  it  likely  that  I  could  have 
overlooked  it  in  so  large  an  insect.  After  examining  Cychrus,  and 
reading  what  Westring  has  written  about  this  beetle,  it  seems  very 
doubtful  whether  it  possesses  any  true  rasp,  though  it  has  the 
power  of  emitting  a  sound. 

From  the  analogy  of  the  Orthoptera  and  Homoptera,  I  expected 
to  find  the  stridulating  organs  in  the  Coleoptera  differing  accord- 
ing to  sex;  but  Landois,  who  has  carefully  examined  several 
species,  observed  no  such  difference;  nor  did  Westring;  nor  did 
Mr.  G.  R.  Crotch  in  preparing  the  many  specimens  which  he  had 
the  kindness  to  send  me.  Any  difference  in  these  organs,  if  slight, 
would,  however,  be  difficult  to  detect,  on  account  of  their  great 
variability.  Thus,  in  the  first  pair  of  specimens  of  Necrophorus 
humator  and  of  Pelobius  which  I  examined,  the  rasp  was  consider- 
ably larger  in  the  male  than  in  the  female;  but  not  so  with  suc- 
ceeding specimens.  In  Geotrupes  stercorarius  the  rasp  appeared 
to  me  thicker,  opaquer,  and  more  prominent  in  three  males  than 
in  the  same  number  of  females;  in  order,  therefore,  to  discover 
whether  the  sexes  differed  in  their  power  of  stridulating,  my  son, 
Mr.  F.  Darwin,  collected  fifty-seven  living  specimens,  which  he 
separated  into  two  lots,  according  as  they  made  a  greater  or  lesser 
noise,  when  held  in  the  same  manner.    He  then  examined  all 


302  THE  DESCENT  OF  MAN. 

these  specimens,  and  found  that  the  males  were  very  nearly  in  the 
same  proportion  to  the  females  in  both  the  lots.  Mr,  F,  Smith  has 
kept  alive  numerous  specimens  of  Monoynchus  pseudacori  (Cur- 
culionidse),  and  is  convinced  that  both  sexes  stridulate,  and  ap- 
parently in  an  equal  degree. 

Nevertheless,  the  power  of  stridulating  is  certainly  a  sexual  char- 
acter in  some  few  Coleoptera.  Mr.  Crotch  discovered  that  the 
males  alone  of  two  species  of  Heliopathes  (Tenebrionidse)  possess 
stridulating  organs.  I  examined  five  males  of  H.  gibbus,  and  in  all 
these  there  was  a  well-developed  rasp,  partially  divided  into  two, 
on  the  dorsal  surface  of  the  terminal  abdominal  segment;  whilst 
in  the  same  number  of  females  there  was  not  even  a  rudiment  of 
the  rasp,  the  membrane  of  this  segment  being  transparent,  and 
much  thinner  than  in  the  male.  In  H.  cribratostriatus  the  male 
has  a  similar  rasp,  excepting  that  it  is  not  partially  divided  into 
two  portions,  and  the  female  is  completely  destitute  of  this  or- 
gan; the  male  in  addition  has  on  the  apical  margins  of  the  elytra, 
on  each  side  of  the  suture,  three  or  four  short  longitudinal  ridges, 
which  are  crossed  by  extremely  fine  ribs,  parallel  to  and  resem- 
bling those  on  the  abdominal  rasp;  whether  these  ridges  serve  as 
an  independent  rasp,  or  as  a  scraper  for  the  abdominal  rasp,  I 
could  not  decide;  the  female  exhibits  no  trace  of  this  latter  struc- 
ture. 

Again,  in  three  species  of  the  Lamellicorn  ^enus  Oryctes,  we 
have  a  nearly  parallel  case.  In  the  females  of  O.  gryphus  and 
nasicornis  the  ribs  on  the  rasp  of  the  pro-pygidium  are  less  con- 
tinuous and  less  distinct  than  in  the  males;  but  the  chief  differ- 
ence is  that  the  whole  upper  surface  of  this  segment,  when  held 
in  the  proper  light,  is  seen  to  be  clothed  with  hairs,  which  are  ab- 
sent or  are  represented  by  excessively  fine  down  in  the  males.  It 
should  be  noticed  that  in  all  Coleoptera  the  effective  part  of  the 
rasp  is  destitute  of  hairs.  In  O.  senegalensis  the  difference  be- 
tween the  sexes  is  more  strongly  marked,  and  this  is  best  seen 
when  the  proper  abdominal  segment  is  cleaned  and  viewed  as  a 
transparent  object.  In  the  female  the  whole  surface  is  covered 
■v\ith  little  separate  crests,  bearing  spines;  whilst  in  the  male 
these  crests  in  proceeding  towards  the  apex,  become  more 
and  more  confluent,  regular,  and  naked;  so  that  three-fourths 
of  the  segment  is  covered  with  extremely  fine  parallel  ribs,  which 
are  quite  absent  in  the  female.  In  the  females,  however,  of  all 
three  species  of  Oryctes,  a  slight  grating  or  stridulating  sound  is 
produced,  when  the  abdomen  of  a  softened  specimen  is  pushed 
backwards  and  forwards. 

In  the  case  of  the  Heliopathes  and  Oryctes  there  can  hardly  be  a 
doubt  that  the  males  stridulate  in  order  to  call  or  to  excite  the 
females;  but  with  most  beetles  the  stridulation  apparently  serves 
both  sexes  as  a  mutual  call.    Beetles  stridulate  under  various  emo- 


COLEOPTERA.  303 

tions,  in  the  same  manner  as  birds  use  their  voices  for  many  pur- 
poses besides  singing  to  their  mates.  The  great  Chiasognathus 
stridulates  in  anger  or  defiance;  many  species  do  the  same  from 
distress  or  fear,  if  held  so  that  they  cannot  escape;  by  striking  the 
hollow  stems  of  trees  in  the  Canary  Islands,  Messrs.  Wollaston 
and  Crotch  were  able  to  discover  the  presence  of  beetles  belonging 
to  the  genus  Acalles  by  their  stridulation.  Lastly,  the  male  Ateu- 
chus  stridulates  to  encourage  the  female  in  her  work,  and  from 
distress  when  she  is  removed. '^^  Some  naturalists  believe  that 
beetles  make  this  noise  to  frighten  away  their  enemies;  but  I  can- 
not think  that  a  quadruped  or  bird,  able  to  devour  a  large  beetle, 
would  be  frightened  by  so  slight  a  sound.  The  belief  that  the 
stridulation  serves  as  a  sexual  call  is  supported  by  the  fact  that 
death-ticks  (Anobium  tessellatum)  are  well  known  to  answer  each 
other's  ticking,  and,  as  I  have  myself  observed,  a  tapping  noise 
artificially  made.  Mr.  Doubleday  also  informs  me  that  he  has 
sometimes  observed  a  female  ticking,^''  and  in  an  hour  or  two  after- 
wards has  found  her  united  with  a  male,  and  on  one  occasion 
surrounded  by  several  males.  Finally,  it  is  probable  that  the 
two  sexes  of  many  kinds  of  beetles  were  at  first  enabled  to  find 
each  other  by  the  slight  shufiling  noise  produced  by  the  rubbing 
together  of  the  adjoining  hard  parts  of  their  bodies;  and  that  as 
those  males  or  females  which  made  the  greatest  noise  succeeded 
best  in  finding  partners,  rugosities  on  various  parts  of  their  bodies 
were  gradually  developed  by  means  of  sexual  selection  into  true 
stridulating  organs. 

79  M.  P.  de  la  Brulerie,  as  quoted  in  'Journal  of  Travel,'  A.  Murray, 
vol.  i.  1868,  p.  135. 

80  According  to  Mr.  Doubleday,  "the  noise  is  produced  by  the  insect 
"raising  itself  on  its  legs  as  high  as  it  can,  and  then  striking  its  thorax 
"five  or  six  times,  in  rapid  succession,  against  the  substance  upon 
"which  it  is  sitting."  For  references  on  this  subject  see  Landois, 
'Zeitschrift  fur  wissen.  Zoolog.'  B.  xvii.  s.  131.  Oliver  says  (as  quoted 
by  Kirby  and  Spence,  'Introduct.'  vol.  ii.  p.  395)  that  the  female  of 
Pimelia  striata  produces  a  rather  loud  sound  by  striking  her  abdomen 
against  any  hard  substance,  "and  that  the  male,  obedient  to  this  call, 
"soon  attends  her,  and  they  pair." 


S04  THE  DESCENT  OF  MAN. 


CHAPTER   XL 

INSECTS,  CONTINUED.— ORDER  LEPIDOPTERA. 

(Butterflies  and  Moths.) 

Courtship  of  butterflies — Battles — Ticking-  noise— Colors  common  to 
both  sexes,  or  more  brilliant  in  the  males— Examples— Not  due  to  the 
direct  action  of  the  conditions  of  life — Colors  adapted  for  protec- 
tion—Colors of  moths— Display— Perceptive  powers  of  the  Lepi- 
doptera — Variability — Causes  of  the  difference  in  color  between  the 
males  and  females— Mimicry,  female  butterflies  more  brilliantly 
colored  than  the  males— Bright  colors  of  caterpillars— Summary  and 
concluding-  remarks  on  the  secondary  sexual  characters  of  insects 
— Birds  and  insects  compared. 

In  this  great  Order  the  most  interesting  points  for  us  are  the 
differences  in  color  between  the  sexes  of  the  same  species,  and  be- 
tween the  distinct  species  of  the  same  genus.  Nearly  the  whole 
of  the  following  chapter  will  he  devoted  to  this  subject;  but  I  will 
first  make  a  few  remarks  on  one  or  two  other  points.  Several 
males  may  often  be  seen  pursuing  and  crowding  round  the  same 
female.  Their  courtship  appears  to  be  a  prolonged  affair,  for  I 
have  frequently  watched  one  or  more  males  pirouetting  round  a 
female  until  I  was  tired,  without  seeing  the  end  of  the  courtship. 
Mr.  A.  G.  Butler  also  informs  me  that  he  has  several  times  watched 
a  male  courting  a  female  for  a  full  quarter  of  an  hour;  but  she 
pertinaciously  refused  him,  and  at  last  settled  on  the  ground  and 
closed  her  wings,  so  as  to  escape  from  his  addresses. 

Although  butterflies  are  weak  and  fragile  creatures,  they  are 
pugnacious,  and  an  Emperor  butterfly^  has  been  captured  with  the 
tips  of  its  wings  broken  from  a  conflict  with  another  male.  Mr. 
Collingwood,  in  speaking  of  the  frequent  battles  between  the  but- 
terflies of  Borneo,  says,  "They  whirl  round  each  other  with  the 
"greatest  rapidity,  and  appear  to  be  incited  by  the  greatest  fe- 
"rocity." 

The  Ageronia  feronia  makes  a  noise  like  that  produced  by  a 

1  Apatura  Iris:    'The  Entomologist's  Weekly  Intelligence,'  1859,  p.  139. 
For    the    Bornean    Butterflies,    see    C.    Collingrwood,    'Rambles    of   a 

Naturalist,'  186S,   p.  183. 


BUTTERFLIES   AND    MOTHS.  305 

toothed  wheel  passing  under  a  spring  catch,  and  which  can  be 
heard  at  the  distance  of  several  yards:  I  noticed  this  sound  at 
Rio  de  Janeiro,  only  when  two  of  these  butterflies  were  chasing 
each  other  in  an  irregular  course,  so  that  it  is  prohahly  made  dur- 
ing the  courtship  of  the  sexes.- 

Some  moths  also  produce  sounds;  for  instance,  the  males  of 
Thecophora  fovea.  On  two  occasions  Mr.  F.  Buchanan  White^ 
heard  a  sharp  quick  noise  made  by  the  male  of  Hylophila  pras- 
inana,  and  which  he  believes  to  be  produced,  as  in  Cicada,  by  an 
elastic  membrane,  furnished  with  a  muscle.  He  quotes,  also, 
Guenee,  that  Setina  produces  a  sound  like  the  ticking  of  a  watch, 
apparently  by  the  aid  of  "two  large  tympaniform  vesicles,  sit- 
"uated  in  the  pectoral  region;"  and  these  "are  much  more 
"developed  in  the  male  than  in  the  female."  Hence  the  sound- 
producing  organs  in  the  Lepidoptera  appear  to  stand  in  some  re- 
lation with  the  sexual  functions.  I  have  not  alluded  to  the  well- 
known  noise  made  by  the  Death's  Head  Sphinx,  for  it  is  generally 
heard  soon  after  the  moth  has  emerged  from  its  cocoon. 

Girard  has  always  observed  that  the  musky  odor,  which  is 
emitted  by  two  species  of  Sphinx  moths,  is  peculiar  to  the  males ;^ 
and  in  the  higher  classes  we  shall  meet  with  many  instances  of 
the  males  alone  being  odoriferous. 

Every  one  must  have  admired  the  extreme  beauty  of  many  but- 
terflies and  of  some  moths;  and  it  may  be  asked,  are  their  colors 
and  diversified  patterns  the  result  of  the  direct  action  of  the 
physical  conditions  to  which  these  insects  have  been  exposed, 
without  any  benefit  being  thus  derived?  Or  have  successive  va- 
riations been  accumulated  and  determined  as  a  protection,  or  for 
some  unknown  purpose,  or  that  one  sex  may  be  attractive  to  the 
other?  And,  again,  what  is  the  meaning  of  the  colors  being 
widely  different  in  the  males  and  females  of  certain  species,  and 
alike  in  the  two  sexes  of  other  species  of  the  same  genus?  Before 
attempting  to  answer  these  questions  a  body  of  facts  must  be 
given. 

With  our  beautiful  English  butterflies,  the  admiral,  peacock, 
and  painted  lady  (Vanessae),  as  well  as  many  others,  the  sexes  are 
alike.  This  is  also  the  case  with  the  magnificent  Heliconidse,  ana 
most  of  the  Danaidae  in  the  tropics.  But  in  certain  other  tropical 
groups,  and  in  some  of  our  English  butterflies,  as  the  purple  em- 

8  See  my  'Journal  of  Researches,'  1845,  p.  33.  Mr.  Doubleda^^  has  de- 
tected ('Proc.  Ent.  Soc'  March  3rd,  1845,  p.  123)  a  peculiar  membranous 
sac  at  the  base  of  the  front  wing-s,  which  is  probably  connected  with 
the  production  of  the  sound.  For  the  case  of  Thecophora,  see  'Zoo- 
logical Record,'  1869,  p.  401.  For  Mr,  Buchanan  White's  observations, 
'The  Scottish  Naturalist,'  July  1872,  p.  214. 

3  'The  Scottish  Naturalist,'  July  1872,  p,  213. 

*  'Zoological  Record,'  1869,  p,  347. 
21 


306  THE  DESCENT  OP  MAN. 

peror,  orange-tip,  &c.  (Apartura  Iris  and  Anttiocharis  carda- 
mines),  the  sexes  differ  either  greatly  or  slightly  in  color.  No  lan- 
guage suffices  to  describe  the  splendor  of  the  males  of  some  tropi- 
cal species.  Even  within  the  same  genus  we  often  find  species 
presenting  extraordinary  differences  between  the  sexes,  whilst 
others  have  their  sexes  closely  alike.  Thus  in  the  South  American 
genus  Epicalia,  Mr.  Bates,  to  whom  I  am  indebted  for  most  of  the 
following  facts,  and  for  looking  over  this  whole  discussion,  in- 
forms me  that  he  knows  twelve  species,  the  two  sexes  of  which 
haunt  the  same  stations  (and  this  is  not  always  the  case  with 
butterflies),  and  which,  therefore,  cannot  have  been  differently  af- 
fected by  external  conditions.^  In  nine  of  these  twelve  species  the 
males  rank  amongst  the  most  brilliant  of  all  butterflies,  and  differ 
so  greatly  from  the  comparatively  plain  females  that  they  were 
formerly  placed  in  distinct  genera.  The  females  of  these  nine 
species  resemble  each  other  in  their  general  type  of  coloration; 
and  they  likewise  resemble  both  sexes  of  the  species  in  several 
allied  genera,  found  in  various  parts  of  the  world.  Hence  we 
may  infer  that  these  nine  species,  and  probably  all  the  others  of 
the  genus,  are  descended  from  an  ancestral  form  which  was  col- 
ored in  nearly  the  same  manner.  In  the  tenth  species  the  female 
still  retains  the  same  general  coloring,  but  the  male  resembles 
her,  so  that  he  is  colored  in  a  much  less  gaudy  and  contrasted 
manner  than  the  males  of  the  previous  species.  In  the  eleventh 
and  twelfth  species,  the  females  depart  from  the  usual  type,  for 
they  are  gaily  decorated  almost  like  the  males,  but  in  a  somewhat 
less  degree.  Hence  in  these  two  latter  species  the  bright  colors 
of  the  males  seem  to  have  been  transferred  to  the  females;  whilst 
in  the  tenth  species  the  male  has  either  retained  or  recovered  the 
plain  colors  of  the  female,  as  well  as  of  the  parent-form  of  the 
genus.  The  sexes  in  these  three  cases  have  thus  been  rendered 
nearly  alike,  though  in  an  opposite  manner.  In  the  allied  genus 
Eubagis,  both  sexes  of  some  of  the  species  are  plain-colored  and 
nearly  alike;  whilst  with  the  greater  number  the  males  are  deco- 
rated with  beautiful  metallic  tints  in  a  diversifled  manner,  and 
differ  much  from  their  females.  The  females  throughout  the 
genus  retain  the  same  general  style  of  coloring,  so  that  they  re- 
semble one  another  much  more  closely  than  they  resemble  their 
own  males. 

In  the  genus  Papilio,  all  the  species  of  the  ^neas  group  are 
remarkable  for  their  conspicuous  and  strongly  contrasted  colors, 
and  they  illustrate  the  frequent  tendency  to  gradation  in  the 
amount  of  difference  between  the  sexes.     In  a  few  species,  for 

5  See  also  Mr.  Bates's  paper  in  'Proc.  Ent.  Soc.  of  Philadelphia,' 
1865,  p.  206.  Also  Mr.  Wallace  on  the  same  subject,  in  regard  to 
Diadema,  in  'Transact.  Entomolog.  Soc.  of  London,'  1869,  p.  278. 


BUTTERFLIES    AND    MOTHS.  307 

Instance  in  P.  ascanius,  the  males  and  females  are  alike;  in  uthers 
the  males  are  either  a  little  brighter,  or  very  much  more  superb 
than  the  females.  The  genus  Junonia,  allied  to  our  Vanessae, 
offers  a  nearly  parallel  case,  for  although  the  sexes  of  most  of 
the  species  resemble  each  other,  and  are  destitute  of  rich  colors, 
j^et  in  certain  species,  as  in  J.  cenone,  the  male  is  rather  more 
bright-colored  than  the  female,  and  in  a  few  (for  instance  J. 
andremiaja)  the  male  is  so  different  from  the  female  that  he 
might  be  mistaken  for  an  entirely  distinct  species. 

Another  striking  case  was  pointed  out  to  me  in  the  British 
Museum  by  Mr.  A.  Butler,  namely,  one  of  the  tropical  American 
Thecl^,  in  which  both  sexes  are  nearly  alike  and  wonderfully 
splendid;  in  another  species  the  male  is  colored  in  a  similarly 
gorgeous  manner,  whilst  the  whole  upper  surface  of  the  female 
is  of  a  dull  uniform  brown.  Our  common  little  English  blue 
butterflies  of  the  genus  Lycsena  illustrate  the  various  differences 
in  color  between  the  sexes,  almost  as  well,  though  not  in  so 
striking  a  manner,  as  the  above  exotic  genera.  In  Lyc^na 
agestis  both  sexes  have  wings  of  a  brown  color,  bordered  with 
small  ocellated  orange  spots,  and  are  thus  alike,  in  L.  cegon 
the  wings  of  the  male  are  of  a  fine  blue,  bordered  with  black; 
whilst  those  of  the  female  are  brown,  with  a  similar  border, 
closely  resembling  the  wings  of  L.  agestis.  Lastly,  in  L.  arion 
both  sexes  are  of  a  blue  color  and  are  very  like,  though  in  the 
female  the  edges  of  the  wings  are  rather  duskier,  with  the  black 
spots  plainer;  and  in  a  bright  blue  Indian  species  both  sexes  are 
still  more  alike. 

I  have  given  the  foregoing  details  in  order  to  show,  in  the  first 
place,  that  when  the  sexes  of  butterflies  differ,  the  male  as  a 
general  rule  is  the  more  beautiful,  and  departs  more  from  the 
usual  type  of  coloring  of  the  group  to  which  the  species  be- 
longs. Hence  in  most  groups  the  females  of  the  several  species 
resemble  each  other  much  more  closely  than  do  the  males.  In 
some  cases,  however,  to  which  I  shall  hereafter  allude,  the  females 
are  colored  more  splendidly  than  the  males.  In  the  second  place, 
these  details  have  been  given  to  bring  clearly  before  the  mind 
that  within  the  same  genus,  the  two  sexes  frequently  present 
every  gradation  from  no  difference  in  color,  to  so  great  a  differ- 
ence that  it  was  long  before  the  two  were  placed  by  entomologists 
in  the  same  genus.  In  the  third  place,  we  have  seen  that  when 
the  sexes  nearly  resemble  each  other,  this  appears  due  either 
to  the  male  having  transferred  his  colors  to  the  female,  or  to  the 
male  having  retained,  or  perhaps  recovered,  the  primordial  col- 
ors of  the  group.  It  also  deserves  notice  that  in  those  groups  in 
which  the  sexes  differ,  the  females  usually  somewhat  resemble 
the  males,  so  that  when  the  males  are  beautiful  to  an  extraor- 
dinary degree,  the  females  almost  invariably  exhibit  some  degree 


SOS  THE  DESCENT  OP  MAN. 

of  beauty.  From  the  many  cases  of  gradation  in  the  amount  of 
difference  between  the  sexes,  and  from  the  prevalence  of  the  same 
general  type  of  coloration  throughout  the  whole  of  the  same 
group,  we  may  conclude  that  the  causes  have  generally  been  the 
same  which  have  determined  the  brilliant  coloring  of  the 
males  alone  of  some  species,  and  of  both  sexes  of  other  species. 

As  so  many  gorgeous  butterflies  inhabit  the  tropics,  it  has 
often  been  supposed  that  they  owe  their  colors  to  the  great  heat 
and  moisture  of  these  zones;  but  Mr.  Bates^  has  shown  by  the 
comparison  of  various  closely-allied  groups  of  insects  from  the 
temperate  and  tropical  regions,  that  this  view  cannot  be  main- 
tained; and  the  evidence  becomes  conclusive  when  brilliantly- 
colored  males  and  plain-colored  females  of  the  same  species 
inhabit  the  same  district,  feed  on  the  same  food,  and  follow  ex- 
actly the  same  habits  of  life.  Even  when  the  sexes  resemble 
each  other,  we  can  hardly  believe  that  their  brilliant  and  beau- 
tifully-arranged colors  are  the  purposeless  result  of  the  nature  of 
the  tissues  and  of  the  action  of  the  surrounding  conditions. 

With  animals  of  all  kinds,  whenever  color  has  been  modified 
for  some  special  purpose,  this  has  been,  as  far  as  we  can  judge, 
either  for  direct  or  indirect  protection,  or  as  an  attraction  be- 
tween the  sexes.  With  many  species  of  butterflies  the  upper  sur- 
faces of  the  wings  are  obscure;  and  this  in  all  probability  leads 
to  their  escaping  observation  and  danger.  But  butterflies  would 
be  particularly  liable  to  be  attacked  by  their  enemies  when  at 
rest;  and  most  kinds  whilst  resting  raise  their  wings  vertically 
over  their  backs,  so  that  the  lower  surface  alone  is  exi^osed  to 
view.  Hence  it  is  this  side  which  is  often  colored  so  as  to  imitate 
the  objects  on  which  these  insects  commonly  rest.  Dr.  Rossler, 
I  believe,  first  noticed  the  similarity  of  the  closed  wings  of  cer- 
tain Vanessse  and  other  butterflies  to  the  bark  of  trees.  Many 
analogous  and  striking  facts  could  be  given.  The  most  interesting 
one  is  that  recorded  by  Mr.  Wallace^  of  a  common  Indian  and 
Sumatran  butterfly  (Kallima),  which  disappears  like  magic  when 
it  settles  on  a  bush;  for  it  hides  its  head  and  antennae  between  its 
closed  wings,  which,  in  form,  color  and  veining,  cannot  be  dis- 
tinguished from  a  withered  leaf  with  its  footstalk.  In  some  other 
cases  the  lower  surfaces  of  the  wings  are  brilliantly  colored,  and 
yet  are  protective;  thus  in  Thecla  rubi  the  wings  when  closed 
are  of  an  emerald  green,  and  resemble  the  young  leaves  of  the 
bramble,  on  which  in  spring  this  butterfly  may  often  be  seen 
seated.    It  is  also  remarkable  that  in  very  many  species  in  which 

6  'The  Naturalist  on  the  Amazons,'  vol.  i.  1863,  p.  19. 

'  See  the  interesting  article  in  the  'Westminster  Review,'  July  1867, 
p.  10.  A  woodcut  of  the  Kallima  is  given  by  Mr.  Wallace  in  'Hard- 
Wicke's  Science  Gossip,'  Sept.  1867,  p.  196. 


BUTTERFLIES    AND    MOTHS.  309 

the  sexes  differ  greatly  in  color  on  their  upper  surface,  the  lower 
surface  is  closely  similar  or  identical  in  both  sexes,  and  serves 
as  a  protection.^ 

Although  the  obscure  tints  both  of  the  upper  and  under 
sides  of  many  butterflies  no  doubt  serve  to  conceal  them,  yet  we 
cannot  extend  this  view  to  the  brilliant  and  conspicuous  colors 
on  the  upper  surface  of  such  species  as  our  admiral  and  peacock 
Vanessae,  our  white  cabbage-butterflies  (Pieris),  or  the  great 
swallow-tail  Papilio  which  haunts  the  open  fens — for  these  but- 
terflies are  thus  rendered  visible  to  every  living  creature.  In 
these  species  both  sexes  are  alike;  but  in  the  common  brim- 
stone butterfly  (Gonepteryx  rhamni),  the  male  is  of  an  intense 
yellow,  whilst  the  female  is  much  paler;  and  in  the  orange-tip 
(Anthocharis  cardamines)  the  males  alone  have  their  v/ings  tipped 
with  bright  orange.  Both  the  males  and  females  in  these  cases 
are  conspicuous,  and  it  is  not  credible  that  their  difference  in 
color  should  stand  in  any  relation  to  ordinary  protection.  Prof. 
Weismann  remarks,^  that  the  female  of  one  of  the  Lycsense  ex- 
pands her  brown  wings  when  she  settles  on  the  ground,  and  is 
then  almost  invisible;  the  male,  on  the  other  hand,  as  if  aware  of 
the  danger  incurred  from  the  bright  blue  of  the  upper  surface  of 
his  wings,  rests  with  them  closed;  and  this  shows  that  the  blue 
color  cannot  be  in  any  way  protective.  Nevertheless,  it  is  probable 
that  conspicuous  colors  are  indirectly  beneficial  to  many  species, 
as  a  warning  that  they  are  unpalatable.  For  in  certain  other 
cases,  beauty  has  been  gained  through  the  imitation  of  other 
beautiful  species,  which  inhabit  the  same  district  and  enjoy  an 
immunity  from  attack  by  being  in  some  way  offensive  to  their 
enemies;  but  then  we  have  to  account  for  the  beauty  of  the 
imitated  species. 

As  Mr.  Walsh  has  remarked  to  me,  the  females  of  our  orange- 
tip  butterfly,  above  referred  to,  and  of  an  American  species 
(Anth.  genutia)  probably  show  us  the  primordial  colors  of  the 
parent-species  of  the  genus;  for  both  sexes  of  four  or  five 
widely-distributed  species  are  colored  in  nearly  the  same  manner. 
As  in  several  previous  cases,  we  may  here  infer  that  it  is  the 
males  of  Anth.  cardamines  and  genutia  which  have  departed 
from  the  usual  type  of  the  genus.  In  the  Anth.  sara  from  Cali- 
fornia, the  orange-tips  to  the  wings  have  been  partially  de- 
veloped in  the  female;  but  they  are  paler  than  in  the  male,  and 
slightly  different  in  some  other  respects.  In  an  allied  Indian 
form,  the  Iphias  glaucippe,  the  orange-tips  are  fully  developed  in 
both  sexes.  In  this  Iphias,  as  pointed  out  to  me  by  Mr.  A.  Butler, 
the  under  surface  of  the  wings  marvelously  resembles  a  pale- 


8  Mr.  G.  Fraser,  in  'Nature,'  April  1871,  p.  489. 

»  'Einfluss  der  Isolirung  auf  die  Artbildung-,'  1872,  p.  58. 


310  THE  DESCENT  OF  MAN. 

colored  leaf;  and  in  our  English  orange-tip,  tlie  under  surface 
vesembles  the  flower-head  of  the  wild  parsley,  on  which  the 
butterfly  often  rests  at  night.^°  The  same  reason  which  compels 
us  to  believe  that  the  lower  surfaces  have  here  been  colored  for 
the  sake  of  protection,  leads  us  to  deny  that  the  wings  have 
been  tipped  with  bright  orange  for  the  same  purpose,  especially 
when  this  character  is  confined  to  the  males. 

Most  Moths  rest  motionless  during  the  whole  or  greater  part 
of  the  day  with  their  wings  depressed;  and  the  whole  upper 
surface  is  often  shaded  and  colored  in  an  admirable  manner,  as 
Mr.  Wallace  has  remarked,  for  escaping  detection.  The  front- 
wings  of  the  Bombycidse  and  Noctuidse,^^  when  at  rest,  generally 
overlap  and  conceal  the  hind-wings;  so  that  the  latter  might  be 
brightly  colored  without  much  risk;  and  they  are  in  fact  often 
thus  colored.  During  flight,  moths  would  often  be  able  to  escape 
from  their  enemies;  nevertheless,  as  the  hind-wings  are  then 
fully  exposed  to  view,  their  bright  colors  must  generally  have 
been  acquired  at  some  little  risk.  But  the  following  fact  shows 
how  cautious  we  ought  to  be  in  drawing  conclusions  on  this  head. 
The  common  Yellow  Under-wings  (Triphsena)  often  fly  about 
during  the  day  or  early  evening,  and  are  then  conspicuous  from 
the  color  of  their  hind-wings.  It  would  naturally  be  thought 
that  this  would  be  a  source  of  danger;  but  Mr.  J.  Jenner  Weir 
believes  that  it  actually  serves  them  as  a  means  of  escape,  for 
birds  strike  at  these  brightly  colored  and  fragile  surfaces,  in- 
stead of  at  the  body.  For  instance,  Mr.  Weir  turned  into  his 
aviary  a  vigorous  specimen  of  Triphasna  pronuba,  which  was  in- 
stantly pursued  by  a  robin;  but  the  bird's  attention  being  caught 
by  the  colored  wings,  the  moth  was  not  captured  until  after 
about  fifty  attempts,  and  small  portions  of  the  wings  were  re- 
peatedly broken  off.  He  tried  the  same  experiment,  in  the  open 
air,  with  a  swallow  and  T.  fimbria;  but  the  large  size  of  this 
moth  probably  interfered  with  its  capture.^^  We  are  thus  remind- 
ed of  a  statement  made  by  Mr.  Wallace,^^  namely,  that  in  the 
Brazilian  forests  and  Malayan  islands,  many  common  and  highly- 
decorated  butterflies  are  weak  flyers,  though  furnished  with  a 
broad  expanse  of  wing;  and  they  "are  often  captured  with  pierced 
"and  broken  wings,  as  if  they  had  been  seized  by  birds,  from 
"which  they  had  escaped:    if  the  wings  had  been  much  smaller 

10  See  the  interesting  observations  by  Mr.  T.  W.  Wood,  'The  Student,* 
Sept.  1868,  p.  81. 

11  Mr.  Wallace  in  'Hardwicke's  Science  Gossip,'  Sept.  1867,  p.  193. 

12  See  also,  on  this  subject,  Mr.  Weir's  paper  in  'Transact,  Ent.  Soc' 
1869,  p.  23. 

13  'Westminster  Review,'  July  1867,  p.  16. 


BUTTERFLIES    AND    MOTHS.  311 

"in  proportion  to  the  body,  it  seerns  probable  that  the  insect 
"would  more  frequently  have  been  struck  cr  pierced  in  a  vital 
"part,  and  thus  the  increased  espanse  of  the  wings  may  have  been 
"indirectly  beneficial." 

Display. — The  bright  colors  of  many  butterflies  and  of  some 
moths  are  specially  arranged  for  display,  so  that  they  may  be 
readily  seen.  During  the  night  colors  are  not  visible,  and 
there  can  be  no  doubt  that  the  nocturnal  moths,  taken  as  a 
body,  are  much  less  gaily  decorated  than  butterflies,  all  of 
which  are  diurnal  in  their  habits.  But  the  moths  of  certain 
families,  such  as  the  Zygaenidse,  several  Sphingidae,  Uraniidse, 
some  Arctiidag  and  SaturniidiB,  fly  about  during  the  day  or 
early  evening,  and  many  of  these  are  extremely  beautiful,  being 
far  brighter  colored  than  the  strictly  nocturnal  kinds.  A  few 
exceptional  cases,  however,  of  bright-colored  nocturnal  species 
have  been  recorded." 

There  is  evidence  of  another  kind  in  regard  to  display.  But- 
terflies, as  before  remarked,  elevate  their  wings  when  at  rest, 
but  whilst  basking  in  the  sunshine  often  alternately  raise  and 
depress  them,  thus  exposing  both  surfaces  to  full  view;  and  al- 
though the  lower  surface  is  often  colored  in  an  obscure  manner 
as  a  protection,  yet  in  many  species  it  is  as  highly  decorated 
as  the  upper  surface,  and  sometimes  in  a  very  different  manner. 
In  some  tropical  species  the  lower  surface  is  even  more  brilliantly 
colored  than  the  upper.^^  In  the  English  fritillaries  (Argynnis) 
the  lower  surface  alone  is  ornamented  with  shining  silver.  Never- 
theless, as  a  general  rule,  the  upper  surface,  which  is  probably 
more  fully  exposed,  is  colored  more  brightly  and  diversely  than 
the  lower.  Hence  the  lower  surface  generally  affords  to  en- 
tomologists the  more  useful  character  for  detecting  the  affinities 
of  the  various  species.  Fritz  Miiller  informs  me  that  three 
species  of  Castnia  are  found  near  his  house  in  S.  Brazil:  of  two 
of  them  the  hind-wings  are  obscure,  and  are  always  covered  by 
the  front-wings  when  these  butterflies  are  at  rest;  but  the  third 
species  has  black  hind-wings,  beautifully  spotted  with  red  and 
white,  and  these  are  fully  expanded  and  displayed  whenever  the 
butterfly  rests.    Other  Ruch  cases  could  be  added. 

If  we  now  turn  to  the  enormous  group  of  moths,  which,  as 

14  For  instance,  Lithosia;  but  Prof.  Westwood  ('Modem  Class,  of 
Insects,'  vol.  ii.  p.  390)  seems  surprised  at  this  case.  On  the  relative 
colors  of  diurnal  and  nocturnal  Lepidoptera,  see  ibid.  pp.  333  and  3S2; 
also  Harris,  'Treatise  on  the  Insects  of  New  England,'  1842,  p.  315. 

^  Such  differences  between  the  upper  and  lower  surfaces  of  the  wing-s 
of  several  species  of  Papilio,  may  be  seen  in  the  beautiful  plates  to 
Mr.  Wallace's  'Memoir  on  the  Papilionidae  of  the  Malayan  Region,' 
in  'Transact.  Linn.  Soc,'  vol.  xxv.  part  i.  1865. 


312  THE  DESCENT  OF  MAN. 

I  hear  from  Mr.  Stainton,  do  not  habitually  expbse  the  under 
surface  of  their  wings  to  full  view,  we  find  this  side  very  rarely 
colored  with  a  brightness  greater  than,  or  even  equal  to,  that 
of  the  upper  side.  Some  exceptions  to  the  rule,  either  real  or 
apparent,  must  be  noticed,  as  the  case  of  Hypopyra/''  Mr.  Trimen 
informs  me  that  in  Guenee's  great  work,  three  moths  are  figured, 
in  which  the  under  surface  is  much  the  more  brilliant.  For 
instance,  in  the  Australian  Gastrophora  the  upper  surface  of  the 
fore-wing  is  pale  grayish-ochreous,  while  the  lower  surface  is 
magnificently  ornamented  by  an  ocellus  of  cobalt-blue,  placed 
in  the  midst  of  a  black  mark,  surrounded  by  orange-yellow,  and 
this  by  bluish-white.  But  the  habits  of  these  three  moths  are 
unknown;  so  that  no  explanation  can  be  given  of  their  unusual 
style  of  coloring.  Mr.  Trimen  also  informs  me  that  the  lower 
surface  of  the  wings  in  certain  other  Geometrse"  and  quadrifid 
Noctuas  are  either  more  variegated  or  more  brightly-colored  than 
the  upper  surface;  but  novae  of  these  species  have  the  habit  of 
"holding  their  wings  quite  erect  over  their  backs,  retaining  them 
"in  this  position  for  a  considerable  time,"  and  thus  exposing  the 
under  surface  to  view.  Other  species,  when  settled  on  the  ground 
or  herbage,  now  and  then  suddenly  and  slightly  lift  up  their 
wings.  Hence  the  lower  surface  of  the  wings  being  brighter 
than  the  upper  surface  in  certain  moths  is  not  so  anomalous  as  it 
at  first  appears.  The  Saturniidse  include  some  of  the  most  beau- 
tiful of  all  moths,  their  wings  being  decorated,  as  in  our  British 
Emperor  moth,  with  fine  ocelli;  and  Mr.  T.  W.  Wood^®  observes 
that  they  resemble  butterflies  in  some  of  their  movements;  "for 
"instance,  in  the  gentle  waving  up  and  down  of  the  wings  as  if 
"for  display,  which  is  more  characteristic  of  diurnal  than  of 
"nocturnal  Lepidoptera." 

It  is  a  singular  fact  that  no  British  moths  which  are  brilliantly 
colored,  and,  as  far  as  I  can  discover,  hardly  any  foreign  species, 
differ  much  in  color  according  to  sex;  though  this  is  the  case  with 
many  brilliant  butterflies.  The  male,  however,  of  one  American 
moth,  the  Saturnia  lo,  is  described  as  having  its  fore-wings  deep 
yellow,  curiously  marked  with  purplish-red  spols;  whilst  the 
wings  of  the  female  are  purple-brown,  marked  with  gray  lines." 
The  British  moths  which  differ  sexually  in  color  are  all  brown, 
or  of  various  dull  yellow  tints,  or  nearly  white.  In  several 
species  the  males  are  much  darker  than  the  females,^"  and  these 

16  See  Mr.  Wormald  on  this  moth:    'Proc.  Ent.  Soc'  March  2nd,  1868. 

"  See  also  an  account  of  the  S.  American  g-enus  Erateina  (one  of  the 
Geometrae)  in  'Transact.  Ent.  Soc'  new  series,  vol.  v.  pi.  xv.  and  xvi. 

18  'Proc.  Ent.  Soc.  of  London,'  July  6,  1868,  p.  xxvii. 

i»  Harris,  'Treatise,'  «S;c.,  edited  by  Flint,  1862,  p.  395. 

20  por  instance,  I  observe  in  my  son's  cabinet  that  the  males  are 
darker  than  the  females  in  the  Lasiocampa  quercus,  Odonestis  pota- 


BUTTERFLIES   AND    MOTHS.  313 

belong  to  groups  which  generally  fly  about  during  the  afternoon. 
On  the  other  hand,  in  many  genera,  as  Mr.  Stainton  informs  me, 
the  males  have  the  hind-wings  whiter  than  those  of  the  female — 
of  which  fact  Agrotis  exclamationis  offers  a  good  instance.  In 
the  Ghost  Moth  (Hepiaius  humuli)  the  difference  is  more  strongly 
marked;  the  males  being  white,  and  the  females  yellow  with 
darker  markings.^^  It  is  probable  that  in  these  cases  the  males 
are  thus  rendered  more  conspicuous,  and  more  easily  seen  by 
the  females  whilst  flying  about  in  the  dusk. 

From  the  several  foregoing  facts  it  is  impossible  to  admit  that 
the  brilliant  colors  of  butterflies,  and  of  some  few  moths,  have 
commonly  been  acquired  for  the  sake  of  protection.  We  have 
seen  that  their  colors  and  elegant  patterns  are  arranged  and  ex- 
hibited as  if  for  display.  Hence  I  am  led  to  believe  that  the 
females  prefer  or  are  most  excited  by  the  more  brilliant  males; 
for  on  any  other  supposition  the  males  would,  as  far  as  we  can 
see,  be  ornamented  to  no  purpose.  We  know  that  ants  and 
certain  Lamellicorn  beetles  are  capable  of  feeling  an  attachment 
for  each  other,  and  that  ants  recognize  their  fellows  after  an 
interval  of  several  months.  Hence  there  is  no  abstract  im- 
probability in  the  Lepidoptera,  which  probably  stand  nearly  or 
quite  as  high  in  the  scale  as  these  insects,  having  sufficient  mental 
capacity  to  admire  bright  colors.  They  certainly  discover  flowers 
by  color.  The  Humming-bird  Sphinx  may  often  be  seen  to 
swoop  down  from  a  distance  on  a  bunch  of  flowers  in  the  midst 
of  green  foliage;  and  I  have  been  assured  by  two  persons 
abroad,  that  these  moths  repeatedly  visit  flowers  painted  on  the 
walls  of  a  room,  and  vainly  endeavor  to  insert  their  proboscis  into 
them.    Fritz  Miiller  informs  me  that  several  kinds  of  butterflies 


toria,  Hypogymn  dispur,  Dasychira  pudibunda,  and  Cycnia  mendica. 
In  this  latter  species  the  difEerence  in  color  between  the  two  sexes  is 
strong-ly  marked;  and  Mr.  Wallace  informs  me  that  we  here  have, 
as  he  believes,  an  instance  of  protective  mimicry  confined  to  one  sex, 
as  will  hereafter  be  more  fully  explained.  The  white  female  of  the 
Cycnia  resembles  the  very  common  Spilosoma  menthrasti,  both  sexes 
of  which  are  white;  and  Mr.  Stainton  observed  that  this  latter  moth 
was  rejected  with  utter  disgust  by  a  whole  brood  of  young  turkeys, 
which  were  fond  of  eating  other  moths;  so  that  if  the  Cycnia  was 
commonly  mistaken  by  British  birds  for  the  Spilosoma,  it  would 
escape  being  devoured,  and  its  white  deceptive  color  would  thus  be 
highly  beneficial. 

21  It  is  remarkable,  that  in  the  Shetland  Islands  the  male  of  this 
moth,  instead  of  differing  widely  from  the  female,  frequently  re- 
sembles her  closely  in  color  (see  Mr.  MacLachlan,  'Transact.  Ent.  Soc' 
vol.  ii.  1866,  p.  459).  Mr.  G.  Eraser  suggests  ('Nature,'  April  1871,  p. 
489)  that  at  the  season  of  the  year  when  the  ghost-moth  appears  in 
these  northern  islands,  the  whiteness  of  the  males  would  not  be  needed 
to  render  them  visible  to  the  females  in  the  twilight  night. 


314  THE  DESCENT  OP  MAN. 

in  S.  Brazil,  show  an  unmistakable  preference  for  certain  colors 
over  others:  he  observed  that  they  very  often  visited  the  brill- 
iant red  flowers  of  five  or  six  genera  of  plants,  but  never  the 
white  or  yellow  flowering  species  of  the  same  and  other  genera, 
growing  in  the  same  garden;  and  I  have  received  other  accounts 
to  the  same  effect.  As  I  hear  from  Mr.  Doubleday,  the  common 
white  butterfly  often  flies  down  to  a  bit  of  paper  on  the  ground, 
no  doubt  mistaking  it  for  one  of  its  own  species.  Mr.  Colling- 
wood-2  in  speaking  of  the  difficulty  in  collecting  certain  butter- 
flies in  the  Malay  Archipelago,  states  that  "a  dead  specimen  pinned 
"upon  a  conspicuous  twig  will  often  arrest  an  insect  of  the 
"same  species  in  its  headlong  flight,  and  bring  it  down  within 
"easy  reach  of  the  net,  especially  if  it  be  of  the  opposite  sex." 

The  courtship  of  butterflies  is,  as  before  remarked,  a  pro- 
longed affair.  The  males  sometimes  fight  together  in  rivalry;  and 
many  may  be  seen  pursuing  or  crowding  round  the  same  female. 
Unless,  then,  the  females  prefer  one  male  to  another,  the  pairing 
must  be  left  to  mere  chance,  and  this  does  not  appear  probable. 
If,  on  the  other  hand,  the  females  habitually,  or  even  occasionally, 
prefer  the  more  beautiful  males,  the  colors  of  the  latter  will 
have  been  rendered  brighter  by  degrees,  and  will  have  been 
transmitted  to  both  sexes  or  to  one  sex,  according  to  the  law 
of  inheritance  which  has  prevailed.  The  process  of  sexual  selec- 
tion will  have  been  much  facilitated,  if  the  conclusion  can  be 
trusted,  arrived  at  from  various  kinds  of  evidence  in  the  supple- 
ment to  the  ninth  chapter;  namely,  that  the  males  of  many 
Lepidoptera,  at  least  in  the  imago  state,  greatly  exceed  the 
females  in  number. 

Some  facts,  however,  are  opposed  to  the  belief  that  female 
butterflies  prefer  the  more  beautiful  males;  thus,  as  I  have 
been  assured  by  several  collectors,  fresh  females  may  frequently 
be  seen  paired  with  battered,  faded,  or  dingy  males;  but  this  is 
a  circumstance  which  could  hardly  fail  often  to  follow  from  the 
males  -emerging  from  their  cocoons  earlier  than  the  females. 
With  moths  of  the  family  of  the  Bombycidse,  the  sexes  pair  im- 
mediately after  assuming  the  imago  state;  for  they  cannot  feed, 
owing  to  the  rudimentary  condition  of  their  mouths.  The  females, 
as  several  entomologists  have  remarked  to  me,  lie  in  an  almost 
torpid  state,  and  appear  not  to  evince  the  least  choice  in  regard 
to  their  partners.  This  is  the  case  with  the  common  silk-motli 
(B.  mori),  as  I  have  been  told  by  some  continental  and  English 
breeders.  Dr.  Wallace,  who  has  had  great  experience  in  breeding 
Bombyx  cynthia,  is  convinced  that  the  females  evince  no  choice 
or  preference.  He  has  kept  above  300  of  these  moths  together, 
and  has  often   found   the  most  vigorous   females   mated   with 

-2  'Rambles  of  a  Naturalist  in  the  Chinese  Seas/  1868,  p.  182. 


BUTTERFLIES   AND    MOTHS.  315 

Stunted  males.  The  reverse  appears  to  occur  seldom;  for,  as  he 
believes,  the  more  vigorous  males  pass  over  the  weakly  females, 
and  are  attracted  by  those  endowed  with  most  vitality.  Neverthe- 
less, the  Bombycidse,  though  obscurely-colored,  are  often  beauti- 
ful to  our  eyes  from  their  elegant  and  mottled  shades. 

I  have  as  yet  only  referred  to  the  species  in  which  the  males 
are  brighter  colored  than  the  females,  and  I  have  attributed 
their  beauty  to  the  females  for  many  generations  having  chosen 
and  paired  with  the  more  attractive  males.  But  converse  cases 
occur,  though  rarely,  in  which  the  females  are  more  brilliant 
than  the  males;  and  here,  as  I  believe,  the  males  have  selected 
the  more  beautiful  females,  and  have  thus  slowly  added  to  their 
beauty.  We  do  not  know  why  in  various  classes  of  animals  the 
males  of  some  few  species  have  selected  the  more  beautiful  fe- 
males instead  of  having  gladly  accepted  any  female,  as  seems  to 
be  the  general  rule  in  the  animal  kingdom;  but  if,  contrary  to 
what  generally  occurs  with  the  Lepidoptera,  the  females  were 
much  more  numerous  than  the  males,  the  latter  would  be  likely 
to  pick  out  the  more  beautiful  females.  Mr.  Butler  showed  me 
several  species  of  Callidryas  in  the  British  Museum,  in  some  of 
which  the  females  equalled,  and  in  others  greatly  surpassed  the 
males  in  beauty;  for  the  females  alone  have  the  borders  of  their 
wings  suffused  with  crimson  and  orange,  and  spotted  with  black. 
The  plainer  males  of  these  species  closely  resemble  each  other, 
showing  that  here  the  females  have  been  modified;  whereas 
in  those  cases,  where  the  males  are  the  more  ornate,  it  is  these 
which  have  been  modified,  the  females  remaining  closely  alike. 

In  England  we  have  some  analogous  cases,  though  not  so 
marked.  The  females  alone  of  two  species  of  Thecla  have  a 
bright-purple  or  orange  patch  on  their  fore-vnngs.  In  Hip- 
parchia  the  sexes  do  not  differ  much;  but  it  is  the  female  of 
H.  janira  which  has  a  conspicuous  light-brown  patch  on  her 
wings;  and  the  females  of  some  of  the  other  species  are  brighter 
colored  than  their  males.  Again,  the  females  of  Colias  edusa  and 
hyale  have  "orange  or  yellow  spots  on  the  black  marginal  border, 
"represented  in  the  males  only  by  thin  streaks;"  and  in  Pieris 
it  is  the  females  which  "are  ornamented  with  black  spots  on  the 
"fore-wings,  and  these  are  only  partially  present  in  the  males." 
Now  the  males  of  many  butterflies  are  known  to  support  the 
females  during  their  marriage  flight;  but  in  the  species  just 
named  it  is  the  females  which  support  the  males;  so  that  the 
part  which  the  two  sexes  play  is  reversed,  as  is  their  relative 
beauty.  Throughout  the  animal  kingdom  the  males  commonly 
take  the  more  active  share  in  wooing,  and  their  beauty  seems 
to  have  been  increased  by  the  females  having  accepted  the  more 
attractive  individuals;  but  with  these  butterflies,  the  females 
take  the  more  active  part  in  the  final  marriage  ceremony,  so 


316  THE  DESCENT  OF  MAN. 

that  we  may  suppose  that  they  likewise  do  so  in  the  wooing;  and 
in  this  case  we  can  understand  hew  it  is  that  they  have  heen 
rendered  the  more  beautiful.  Mr.  Meldola,  from  whom  the  fore- 
going statements  have  been  taken,  says  in  conclusion:  "Though 
"I  am  not  convinced  of  the  action  of  sexual  selection  in  produc- 
"ing  the  colors  of  insects,  it  cannot  be  denied  that  these  facts  are 
"strikingly  corroborative  of  Mr.  Darwin's  views." 


"23 


As  sexual  selection  primarily  depends  on  variability,  a  few 
words  must  be  added  on  this  subject.  In  respect  to  color  there 
is  no  difficulty,  for  any  number  of  highly  variable  Lepidoptera 
could  be  named.  One  good  instance  will  suffice.  Mr.  Bates 
showed  me  a  whole  series  of  specimens  of  Papilio  sesostris  and 
P.  childrense;  in  the  latter  the  males  varied  much  in  the  extent 
of  the  beautifully  enamelled  green  patch  on  the  fore-wings,  and 
in  the  size  of  the  white  mark,  and  of  the  splendid  crimson  stripe 
on  the  hind-wings;  so  that  there  was  a  great  contrast  amongst 
the  males  between  the  most  and  the  least  gaudy.  The  male  of 
Papilio  sesostris  is  much  less  beautiful  than  of  P.  childrense; 
and  it  likewise  varies  a  little  in  the  size  of  the  green  patch  on  the 
fore-wings,  and  in  the  occasional  appearance  of  the  small  crimson 
stripe  on  the  hind-wings,  borrowed,  as  it  would  seem,  from  its 
own  female;  for  the  females  of  this  and  of  many  other  species 
in  the  ^neas  group  possess  this  crimson  stripe.  Hence  between 
the  brightest  specimens  of  P.  sesostris  and  the  dullest  of  P.  chil- 
drense, there  was  but  a  small  interval;  and  it  was  evident  that  as 
far  as  mere  variability  is  concerned,  there  would  be  no  difficulty 
in  permanently  increasing  the  beauty  of  either  species  by  means 
of  selection.  The  variability  is  here  almost  confined  to  the  male 
sex;  but  Mr.  Wallace  and  Mr.  Bates  have  shown-*  that  the  females 
of  some  species  are  extremely  variable,  the  males  being  nearly 
constant.  In  a  future  chapter  I  shall  have  occasion  to  show  that 
the  beautiful  eye-like  spots,  or  ocelli,  found  on  the  wings  of 
many  Lepidoptera,  are  eminently  variable.  I  may  here  add  that 
these  ocelli  offer  a  difficulty  on  the  theory  of  sexual  selection; 
for  though  appearing  to  us  so  ornamental,  they  are  never  pres- 
ent in  one  sex  and  absent  in  the  other,  nor  do  they  ever  differ 


23  'Nature,'  April  27th,  1871,  p.  508.  Mr.  Meldola  quotes  Donzel,  in 
'Soc.  Ent.  de  France,'  1837,  p.  77,  on  the  flight  of  butterflies  whilst 
pairing-.  See  also  Mr.  G.  Fraser,  in  'Nature,'  April  20th,  1871,  p.  489,  on 
the  sexual  differences  of  several  British  butterflies. 

24  Wallace  on  the  Papilionidae  of  the  Malayan  Region,  in  'Transact. 
Linn.  Soc'  vol.  xxv.  1865,  pp.  8,  36.  A  striking  case  of  a  rare  variety, 
strictly  intermediate  between  two  other  well-marked  female  varieties, 
is  given  by  Mr.  "Wallace.  See  also  Mr.  Bates,  in  'Proc.  Entomologr. 
Soc'  Nov.  19th,  1866,  p.  xl. 


BUTTERFLIES    AND    MOTHS.  317 

much  in  the  two  sexes.-"  This  fact  is  at  present  inexplicable; 
but  if  it  should  hereafter  be  found  that  the  formation  of  an  ocellus 
is  due  to  some  change  in  the  tissues  of  the  wings,  for  instance, 
occurring  at  a  very  early  period  of  development,  we  might  expect, 
from  what  we  know  of  the  laws  of  inheritance,  that  it  would  be 
transmitted  to  both  sexes,  though  arising  and  perfected  in  one 
sex  alone. 

On  the  whole,  though  many  serious  objections  may  be  urged, 
it  seems  probable  that  most  of  the  brilliantly  colored  species  of 
Lepidoptera  owe  their  colors  to  sexual  selection,  excepting  in 
certain  cases,  presently  to  be  mentioned,  in  which  conspicuous 
colors  have  been  gained  through  mimicry  as  a  protection.  From 
the  ardor  of  the  male  throughout  the  animal  kingdom,  he  is 
generally  willing  to  accept  any  female;  and  it  is  the  female  which 
usually  exerts  a  choice.  Hence,  if  sexual  selection  has  been 
efficient  with  the  Lepidoptera,  the  male,  when  the  sexes  differ, 
ought  to  be  the  more  brilliantly  colored,  and  this  undoubtedly  is 
the  case.  When  both  sexes  are  brilliantly  colored  and  resemble 
each  other,  the  characters  acquired  by  the  males  appear  to  have 
been  transmitted  to  both.  We  are  led  to  this  conclusion  by  cases, 
even  within  the  same  genus,  of  gradation  from  an  extraordinary 
amount  of  difference  to  identity  in  color  between  the  two  sexes. 

But  it  may  be  asked  whether  the  differences  in  color  between 
the  sexes  may  not  be  accounted  for  by  other  means  besides 
sexual  selection.  Thus  the  males  and  females  of  the  same  species 
of  butterfxy  are  in  several  cases  known^^  to  inhabit  different  sta- 
tions, the  former  commonly  basking  in  the  sunshine,  the  latter 
haunting  gloomy  forests.  It  is  therefore  possible  that  different 
conditions  of  life  may  have  acted  directly  on  the  two  sexes;  but 
this  is  not  probable,"  as  in  the  adult  state  they  are  exposed  to 
different  conditions  during  a  very  short  period;  and  the  larvae 
of  both  are  exposed  to  the  same  conditions.  Mr.  Wallace  be- 
lieves that  the  difference  between  the  sexes  is  due  not  so  much 
to  the  males  having  been  modified,  as  to  the  females  having  in  all 
or  almost  all  cases  acquired  dull  colors  for  the  sake  of  protection. 
It  seems  to  me,  on  the  contrary,  far  more  probable  that  it  is  the 
males  which  have  been  chiefly  modified  through  sexual  selec- 
tion, the  females  having  been  comparatively  little  changed. 
We  can  thus  understand  how  it  is  that  the  females  of  allied 
species  generally  resemble  one  another  so  much  more  closely 

25  Mr.  Bates  was  so  kind  as  to  lay  this  subject  before  the  Entomo- 
logical Society,  and  I  have  received  answers  to  this  effect  from  sev- 
eral entomologists. 

28  H.  W.  Bates,  'The  Naturalist  on  the  Amazons,'  vol.  ii.  1863,  p.  228. 
A.  R.  Wallace,  in  'Transact.  Linn.  SoC  vol.  xxv.  1865,  p.  10. 

27  On  this  whole  subject  see  'The  Variation  of  Animals  and  Plants 
under  Domestication,'  1868,  vol.  ii.  chap,  xxiii. 


313  THE  DESCENT  OF  MAN. 

than  do  the  males.  They  thus  show  us  approximately  the  pri- 
mordial coloring  of  the  parent-species  of  the  group  to  which  they 
belong.  They  have,  however,  almost  always  been  somewhat  modi- 
fied by  the  transfer  to  them  of  some  of  the  successive  variations, 
through  the  accumulation  of  which  the  males  were  rendered  beau- 
tiful. But  I  do  not  wish  to  deny  that  the  females  alone  of  some 
species  may  have  been  specially  modified  for  protection.  In  most 
cases  the  males  and  females  of  distinct  species  will  have  been 
exposed  during  their  prolonged  larval  state  to  different  conditions, 
and  may  have  been  thus  affected;  though  with  the  males  any 
slight  change  of  color  thus  caused  will  generally  have  been 
masked  by  the  brilliant  tints  gained  through  sexual  selection. 
When  we  treat  of  Birds,  I  shall  have  to  discuss  the  whole  question, 
as  to  how  far  the  differences  in  color  between  the  sexes  are  due 
to  the  males  having  been  modified  through  sexual  selection  for 
ornamental  purposes,  or  to  the  females  having  been  modified 
through  natural  selection  for  the  sake  of  protection,  so  that  I  will 
here  say  but  little  on  the  subject. 

In  all  the  cases  in  which  the  more  common  form  of  equal 
inheritance  by  both  sexes  has  prevailed,  the  selection  of  bright- 
colored  males  would  tend  to  make  the  females  bright-colored; 
and  the  selection  of  dull-colored  females  would  tend  to  make 
the  males  dull.  If  both  processes  were  carried  on  simultaneously, 
they  would  tend  to  counteract  each  other;  and  the  final  result 
would  depend  on  whether  a  greater  number  of  females  from 
being  well  protected  by  obscure  colors,  or  a  greater  number  of 
males  by  being  brightly-colored  and  thus  finding  partners,  suc- 
ceeded in  leaving  more  numerous  offspring. 

In  order  to  account  for  the  frequent  transmission  of  characters 
to  one  sex  alone,  Mr.  Wallace  expresses  his  belief  that  the  more 
common  form  of  equal  inheritance  by  both  sexes  can  be  changed 
through  natural  selection  into  inheritance  by  one  sex  alone,  but 
in  favor  of  this  view  I  can  discover  no  evidence.  We  know  from 
what  occurs  under  domestication  that  new  characters  often  ap- 
pear, which  from  the  first  are  transmitted  to  one  sex  alone;  and 
by  the  selection  of  such  variations  there  v/ould  not  be  the  slight- 
est difficulty  in  giving  bright  colors  to  the  males  alone,  and  at 
the  same  time  or  subsequently,  dull  colors  to  the  females  alone. 
In  this  manner  the  females  of  some  butterflies  and  moths  have,  it 
is  probable,  been  rendered  inconspicuous  for  the  sake  of  pro- 
tection, and  widely  different  from  their  males. 

I  am,  however,  unwilling  without  distinct  evidence  to  admit 
that  two  complex  processes  of  selection,  each  requiring  the 
transference  of  new  characters  to  one  sex  alone,  have  been  carried 
on  with  a  multitude  of  species, — that  the  males  have  been  rendered 
more  brilliant  by  beating  their  rivals,  and  the  females  more  dull- 
colored  by  having  escaped  from  their  enemies.    The  male,  for 


BUTTERFLIES    AND    MOTHS.  319 

instance,  of  the  comfnon  brimstone  butterfly  (Gonepteryx),  is  of 
a  far  more  intense  yellow  than  the  female,  though  she  is  equally 
conspicuous;  and  it  does  not  seem  probable  that  she  specially 
acquired  her  pale  tints  as  a  protection,  though  it  is  probable 
that  the  male  acquired  his  bright  colors  as  a  sexual  attraction. 
The  female  of  Anthocharis  cardamines  does  not  possess  the 
beautiful  orange  wing-tips  of  the  male;  consequently  she  closely 
resembles  the  white  butterflies  (Pieris)  so  common  in  our  gar- 
dens; but  we  have  no  evidence  that  this  resemblance  is  bene- 
ficial to  her.  As,  on  the  other  hand,  she  resembles  both  sexes 
of  several  other  species  of  the  genus  inhabiting  various  quarters 
of  the  world,  it  is  probable  that  she  has  simply  retained  to  a 
large  extent  her  primordial  colors. 

Finally,  as  we  have  seen,  various  considerations  lead  to  the 
conclusion  that  with  the  greater  number  of  brilliantly-colored 
Lepidoptera  it  is  the  male  which  has  been  chiefly  modified 
through  sexual  selection;  the  amount  of  difference  between  the 
sexes  mostly  depending  on  the  form  of  inheritance  which  has 
prevailed.  Inheritance  is  governed  by  so  many  unknown  laws 
or  conditions,  that  it  seems  to  us  to  act  in  a  capricious  manner;^** 
and  we  can  thus,  to  a  certain  extent,  understand  how  it  is  that 
with  closely  allied  species  the  sexes  either  differ  to  an  astonish- 
ing degree,  or  are  identical  in  color.  As  all  the  successive  steps 
in  the  process  of  variation  are  necessarily  transmitted  through 
the  female,  a  greater  or  less  number  of  such  steps  might  readily 
become  developed  in  her;  and  thus  we  can  understand  the  fre- 
quent gradations  from  an  extreme  difference  to  none  at  all  be- 
tween the  sexes  of  allied  species.  These  cases  of  gradation,  it 
may  be  added,  are  much  too  common  to  favor  the  supposition 
that  we  here  see  females  actually  undergoing  the  process  of 
transition  and  losing  their  brightness  for  the  sake  of  protection; 
for  we  have  every  reason  to  conclude  that  at  any  one  time  the 
greater  number  of  species  are  in  a  fixed  condition. 

Mimicry. — This  principle  was  first  made  clear  in  an  admir- 
able paper  by  Mr.  Bates,-^  who  thus  threw  a  flood  of  light  on 
many  obscure  problems.  It  had  previously  been  observed  that 
certain  butterflies  in  S.  America  belonging  to  quite  distinct  fam- 
ilies, resembled  the  Heliconidse  so  closely  in  every  stripe  and 
shade  of  color,  that  they  could  not  be  distinguished  save  by  an 
experienced  entomologist.  As  the  Heliconidae  are  colored  in 
their  usual  manner,  whilst  the  others  depart  from  the  usual  color- 
ing of  the  groups  to  which  they  belong,  it  is  clear  that  the  latter 

28  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
ii.  chap.  xii.  p.  17. 

29  -Transact.  Linn.  Soc'  vol.  xxiii.  1862,  p.  495. 


320  THE  DESCENT  OF  MAN. 

are  the  imitators,  and  the  Heliconidse  the  imitated.  Mr.  Bates 
further  observed  that  the  imitating  species  are  comparatively 
rare,  whilst  the  imitated  abound,  and  that  the  two  sets  live 
mingled  together.  From  the  fact  of  the  Heliconidae  being  con- 
spicuous and  beautiful  insects,  yet  so  numerous  in  individuals 
and  species,  he  concluded  that  they  must  be  protected  from  the 
attacks  of  enemies  by  some  secretion  or  odor;  and  this  conclusion 
has  now  been  amply  confirmed,""  especially  by  Mr.  Belt.  Hence 
Mr.  Bates  inferred  that  the  butterflies  which  imitate  the  protected 
species  have  acquired  their  present  marvelously  deceptive  ap- 
pearance through  variation  and  natural  selection,  in  order  to  be 
mistaken  for  the  protected  kinds,  and  thus  to  escape  being  de- 
voured. No  explanation  is  here  attempted  of  the  brilliant  colors 
of  the  imitated,  but  only  of  the  imitating  butterflies.  We  must 
account  for  the  colors  of  the  former  in  the  same  general  manner, 
as  in  the  cases  previously  discussed  in  this  chapter.  Since  the 
publication  of  Mr.  Bates'  paper,  similar  and  equally  striking 
facts  have  been  observed  by  Mr.  Wallace  in  the  Malayan  region, 
by  Mr.  Trimen  in  South  Africa,  and  by  Mr.  Riley  in  the  United 
States.^^ 

As  some  writers  have  felt  much  difficulty  in  understanding 
how  the  first  steps  in  the  process  of  mimicry  could  have  been 
effected  through  natural  selection,  it  may  be  well  to  remark  that 
the  process  probably  commenced  long  ago  between  forms  not 
widely  dissimilar  in  color.  In  this  case  even  a  slight  variation 
would  be  beneficial,  if  it  rendered  the  one  species  more  like  the 
other;  and  afterwards  the  imitated  species  might  b©  modified  to 
an  extreme  degree  through  sexual  selection  or  other  means,  and  if 
the  changes  were  gradual,  the  imitators  might  easily  be  led  along 
the  same  track,  until  they  differed  to  an  equally  extreme  degree 
from  their  original  condition;  and  they  would  thus  ultimately 
assume  an  appearance  or  coloring  wholly  unlike  that  of  the 
other  members  of  the  group  to  which  they  belonged.  It  should 
also  be  remembered  that  many  species  of  Lepidoptera  are  liable 
to  considerable  and  abrupt  variations  in  color.  A  few  instances 
have  been  given  in  this  chapter;  and  many  more  may  be  found 
in  the  papers  of  Mr.  Bates  and  Mr.  Wallace. 

With  several  species  the  sexes  are  alike,  and  imitate  the  two 
sexes  of  another  species.  But  Mr.  Trimen  gives,  in  the  paper 
already  referred  to,  three  cases  in  which  the  sexes  of  the  imitated 

30  'Proc.  Ent.  Soc'  Dec.  3rd,  1866,  p.  xlv. 

31  Wallace,  'Transact.  Linn.  Soc'  vol.  xxv.  1865,  p.  1;  alsio  'Transact. 
Ent.  Soc'  vol.  iv.  (3rd  series),  1867,  p.  301.  Trimen,  'Linn.  Transact.' 
vol.  xxvi.  1869,  p.  497.  Riley,  'Third  Annual  Report  on  the  Noxious 
Insects  of  Missouri,'  1871,  pp.  163-168.  This  latter  essay  is  valuable,  as 
Mr.  Riley  here  discusses  all  the  objections  which  have  been  raised 
against  Mr.  Bates*  theory. 


BUTTERFLIES   AND    MOTHS.  321 

form  differ  from  each  other  in  color,  and  the  sexes  of  the  imitat- 
ing form  differ  in  a  like  manner.  Several  cases  have  also  been 
recorded  where  the  females  alone  imitate  brilliantly-colored  and 
protected  species,  the  males  retaining  "the  normal  aspect  of  their 
"immediate  congeners."  It  is  here  obvious  that  the  successive 
variations  by  which  the  female  has  been  modified  have  been 
transmitted  to  her  alone.  It  is,  however,  probable  that  some 
of  the  many  successive  variations  would  have  been  transmitted 
to,  and  developed  in,  the  males  had  not  such  males  been  elim- 
inated by  being  thus  rendered  less  attractive  to  the  females;  so 
that  only  those  variations  were  preserved  which  were  from  the 
first  strictly  limited  in  their  transmission  to  the  female  sex.  We 
have  a  partial  illustration  of  these  remarks  in  a  statement  by  Mr. 
Belt;^^  that  the  males  of  some  of  the  Leptalides,  which  imitate 
protected  species,  still  retain  in  a  concealed  manner  some  of 
their  original  characters.  Thus  in  the  males  "the  upper  half  of 
"the  lower  wing  is  of  pure  white,  whilst  all  the  rest  of  the  wings 
"is  barred  and  spotted  with  black,  red  and  yellow,  like  the 
"species  they  mimic.  The  females  have  not  this  white  patch,  and 
"the  males  usually  conceal  it  by  covering  it  with  the  upper  wing, 
"so  that  I  cannot  imagine  its  being  of  any  other  use  to  them 
"than  as  an  attraction  in  courtship,  when  they  exhibit  it  to  the 
"females,  and  thus  gratify  their  deep-seated  preference  for  the 
"normal  color  of  the  Order  to  which  the  Leptalides  belong." 

Bright  Colors  of  Caterpillars. — Whilst  reflecting  on  the  beauty 
of  many  butterflies,  it  occurred  to  me  that  some  caterpillars  were 
splendidly  colored;  and  as  sexual  selection  could  not  possibly 
have  here  acted,  it  appeared  rash  to  attribute  the  beauty  of  the 
mature  insect  to  this  agency,  unless  the  bright  colors  of  their 
larvae  could  be  somehow  explained.  In  the  first  place,  it  may  be 
observed  that  the  colors  of  caterpillars  do  not  stand  in  any  close 
correlation  with  those  of  the  mature  insect.  Secondly,  their 
bright  colors  do  not  serve  in  any  ordinary  manner  as  a  protection. 
Mr.  Bates  informs  me,  as  an  instance  of  this,  that  the  most  con- 
spicuous caterpillar  which  he  ever  beheld  (that  of  a  Sphinx)  lived 
on  the  large  green  leaves  of  a  tree  on  the  open  llanos  of  South 
America;  it  was  about  four  inches  in  length,  transversely  banded 
with  black  and  yellow,  and  with  its  head,  legs,  and  tail  of  a 
bright  red.  Hence  it  caught  the  eye  of  any  one  who  passed  by, 
even  at  a  distance  of  many  yards,  and  no  doubt  that  of  every 
passing  bird. 

I  then  applied  to  Mr.  Wallace,  who  has  an  innate  genius  for 
solving  difllculties.  After  some  consideration  he  replied:  "Most 
"caterpillars  require  protection,  as  may  be  inferred  from  some 


'The  Naturalist  in  Nicaragua,'  1874,  p.  385. 
22 


322  THE  DESCENT  OF  MAN. 

"kinds  being  furnished  with  spines  or  irritating  hairs,  and  from 
"many  being  colored  green  like  the  leaves  on  which  they  feed, 
"or  being  curiously  like  the  twigs  of  the  trees  on  which  they 
"live."  Another  instance  of  protection,  furnished  me  by  Mr.  J. 
Mansel  Weale,  may  be  added,  namely,  that  there  is  a  caterpillar 
of  a  moth  which  lives  on  the  mimosas  in  South  Africa,  and 
fabricates  for  itself  a  case  quite  indistinguishable  from  the  sur- 
rounding thorns.  From  such  considerations  Mr.  Wallace  thought 
it  probable  that  conspicuously-colored  caterpillars  were  protected 
by  having  a  nauseous  taste;  but  as  their  skin  is  extremely  ten- 
der, and  as  their  intestines  readily  protrude  from  a  wound,  a 
slight  peck  from  the  beak  of  a  bird  would  be  as  fatal  to  them 
as  if  they  had  been  devoured.  Hence,  as  Mr.  Wallace  remarks, 
"distastefulness  alone  would  be  insuflBcient  to  protect  a  cater- 
"pillar  unless  some  outward  sign  indicated  to  its  would-be  de- 
"stroyer  that  its  prey  was  a  disgusting  morsel."  Under  these 
circumstances  it  would  be  highly  advantageous  to  a  caterpillar 
to  be  instantaneously  and  certainly  recognized  as  unpalatable  by 
all  birds  and  other  animals.  Thus  the  most  gaudy  colors  would 
be  serviceable,  and  might  have  been  gained  by  variation  and  the 
survival  of  the  most  easily-recognized  individuals. 

This  hypothesis  appears  at  first  sight  very  bold,  but  when  it 
was  brought  before  the  Entomological  Society^^  it  was  supported 
by  various  statements;  and  Mr.  J.  Jenner  Weir,  who  keeps  a 
large  number  of  birds  in  an  aviary,  informs  me  that  he  has 
made  many  trials,  and  finds  no  exception  to  the  rule,  that  all 
caterpillars  of  nocturnal  and  retiring  habits  with  smooth  skins, 
all  of  a  green  color,  and  all  which  imitate  twigs,  are  greedily 
devoured  by  his  birds.  The  hairy  and  spinose  kinds  are  invari- 
ably rejected,  as  were  four  conspicuously-colored  species.  When 
the  birds  rejected  a  caterpillar,  they  plainly  showed,  by  shaking 
their  heads,  and  cleansing  their  beaks,  that  they  were  disgusted 
by  the  taste.^*  Three  conspicuous  kinds  of  caterpillars  and  moths 
were  also  given  to  some  lizards  and  frogs,  by  Mr.  A.  Butler,  and 
were  rejected,  though  other  kinds  were  eagerly  eaten.  Thus  the 
probability  of  Mr.  Wallace's  view  is  confirmed,  namely,  that  cer- 
tain caterpillars  have  been  made  conspicuous  for  their  own  good, 
so  as  to  be  easily  recognized  by  their  enemies,  on  nearly  the 
same  principle  that  poisons  are  sold  in  colored  bottles  by  drug- 

33  'Proc.  Entomolog-.  Soc'  Dec.  3rd,  1866,  p.  xlv.,  and  March  4th,  1867, 
p.  Ixxx. 

3*  See  Mr.  J.  Jenner  Weir's  paper  on  Insects  and  Insectivorous  Birds, 
in  'Transact.  Ent.  Soc'  1869,  p.  21,  also  Mr.  Butler's  paper,  ibid.  p.  27. 
Mr.  Riley  has  given  analogous  facts  in  the  'Third  Annual  Report  on 
the  Noxious  Insects  of  Missouri,'  1871,  p.  148.  Some  opposed  cases  are, 
however,  given  by  Dr.  Wallace  and  M.  H.  d'Orville;  see  'Zoological 
Record,'  1869,  p.  349. 


BUTTERFLIES   AND   MOTHS,  323 

gists  for  the  good  of  man.  We  cannot,  however,  at  present  thus 
explain  the  elegant  diversity  in  the  colors  of  many  caterpillars; 
but  any  species  which  had  at  some  former  period  acquired  a  dull, 
mottled,  or  striped  appearance,  either  in  imitation  of  surround- 
ing objects,  or  from  the  direct  action  of  climate,  &c.,  almost  cer- 
tainly would  not  become  uniform  in  color,  when  its  tints  were 
rendered  intense  and  bright;  for  in  order  to  make  a  caterpillar 
merely  conspicuous,  there  would  be  no  selection  in  any  definite 
direction. 

Summainj  and  Concluding  RemarTis  on  Insects. — Looking  back 
to  the  several  Orders,  we  see  that  the  sexes  often  differ  in 
various  characters,  the  meaning  of  which  is  not  in  the  least 
understood.  The  sexes,  also,  often  differ  in  their  organs  of  sense 
and  means  of  locomotion,  so  that  the  males  may  quickly  dis- 
cover and  reach  the  females.  They  differ  still  oftener  in  the 
males  possessing  diversified*  contrivances  for  retaining  the  fe- 
males when  found.  We  are,  however,  here  concerned  only  in  a 
secondary  degree  with  sexual  differences  of  these  kinds. 

In  almost  all  the  Orders,  the  males  of  some  species,  even  of 
weak  and  delicate  kinds,  are  known  to  be  highly  pugnacious; 
and  some  few  are  furnished  with  special  weapons  for  fighting 
with  their  rivals.  But  the  law  of  battle  does  not  prevail  nearly 
so  widely  with  insects  as  with  the  higher  animals.  Hence  it 
probably  arises,  that  it  is  in  only  a  few  cases  that  the  males  have 
been  rendered  larger  and  stronger  than  the  females.  On  the 
contrary,  they  are  usually  smaller,  so  that  they  may  be  developed 
within  a  shorter  time,  to  be  ready  in  large  numbers  for  the 
emergence  of  the  females. 

In  two  families  of  the  Homoptera  and  in  three  of  the  Orthop- 
tera,  the  males  alone  possess  sound-producing  organs  in  an 
efficient  state.  These  are  used  incessantly  during  the  breeding- 
season,  not  only  for  calling  the  females,  but  apparently  for 
charming  or  exciting  them  in  rivalry  with  other  males.  No 
one  who  admits  the  agency  of  selection  of  any  kind,  will,  after 
reading  the  above  discussion,  dispute  that  these  musical  instru- 
ments have  been  acquired  through  sexual  selection.  In  four 
other  Orders  the  members  of  one  sex,  or  more  commonly  of  both 
sexes,  are  provided  with  organs  for  producing  various  sounds, 
which  apparently  serve  merely  as  call-notes.  When  both  sexes 
are  thus  provided,  the  individuals  which  were  able  to  make  the 
loudest  or  most  continuous  noise  would  gain  partners  before 
those  which  were  less  noisy,  so  that  their  organs  have  probably 
been  gained  through  sexual  selection.  It  is  instructive  to  reflect 
on  the  wonderful  diversity  of  the  means  for  producing  sound, 
possessed  by  the  males  alone,  or  by  both  sexes,  in  no  less  than 
six  Orders.     We  thus  learn  how  e^ectual  sexual  selection  has 


324  THE  DESCENT  OF  MAN. 

been  in  leading  to  modifications  which  sometimes,  as  with  the 
Homoptera,  relate  to  important  parts  of  the  organization. 

From  the  reasons  assigned  in  the  last  chapter,  it  is  probable 
that  the  great  horns  possessed  by  the  males  of  many  Lamelli- 
corn,  and  some  other  beetles,  have  been  acquired  as  ornaments. 
From  the  small  size  of  insects,  we  are  apt  to  undervalue  their 
appearance.  If  we  could  imagine  a  male  Chalcosoma  (fig.  16), 
with  its  polished  bronzed  coat  of  mail,  and  its  vast  complex 
horns,  magnified  to  the  size  of  a  horse,  or  even  of  a  dog,  it  would 
be  one  of  the  most  imposing  animals  in  the  world. 

The  coloring  of  insects  is  a  complex  and  obscure  subject. 
When  the  male  differs  slightly  from  the  female,  and  neither  are 
brilliantly-colored,  it  is  probable  that  the  sexes  have  varied 
in  a  slightly  different  manner,  and  that  the  variations  have  been 
transmitted  by  each  sex  to  the  same,  without  any  benefit  or  evil 
thus  accruing.  When  the  male  is  brilliantly-colored  and  differs 
conspicuously  from  the  female,  as  .with  some  dragon-fiies  and 
many  butterflies,  it  is  probable  that  he  owes  his  colors  to  sexual 
selection;  whilst  the  female  has  retained  a  primordial  or  very 
ancient  type  of  coloring,  slightly  modified  by  the  agencies  before 
explained.  But  in  some  cases  the  female  has  apparently  been 
made  obscure  by  variations  transmitted  to  her  alone,  as  a  means 
of  direct  protection;  and  it  is  almost  certain  that  she  has  some- 
times been  made  brilliant,  so  as  to  imitate  other  protected  spe- 
cies inhabiting  the  same  district.  When  the  sexes  resemble  each 
other  and  both  are  obscurely  colored,  there  is  no  doubt  that  they 
have  been  in  a  multitude  of  cases  so  colored  for  the  sake  of  pro- 
tection. So  it  is  in  some  instances  when  both  are  brightly-col- 
ored, for  they  thus  imitate  protected  species,  or  resetnble  sur- 
rounding objects  such  as  flowers;  or  they  give  notice  to  their 
enemies  that  they  are  unpalatable.  In  other  cases  in  which  the 
sexes  resemble  each  other  and  are  both  brilliant,  especially  when 
the  colors  are  arranged  for  display,  we  may  conclude  that  they 
have  been  gained  by  the  male  sex  as  an  attraction,  and  have  been 
transferred  to  the  female.  We  are  more  especially  led  to  this 
conclusion  whenever  the  same  type  of  coloration  prevails  through- 
out a  whole  group,  and  we  find  that  the  males  of  some  species 
differ  widely  in  color  from  the  females,  whilst  others  differ 
slightly  or  not  at  all,  with  intermediate  gradations  connecting 
these  extreme  states. 

In  the  same  manner  as  bright  colors  have  often  been  partially 
transferred  from  the  males  to  the  females,  so  it  has  been  with 
the  extraordinary  horns  of  many  Lamellicorn  and  some  other 
beetles.  So  again,  the  sound-producing  organs  proper  to  the 
males  of  the  Homoptera  and  Orthoptera  have  generally  been 
transferred  in  a  rudimentary,  or  even  in  a  nearly  perfect  condi- 
tion, to  the  females;    yet  not  sufficiently  perfect  to  be  of  any  use. 


SUMMARY    ON    INSECTS.  325 

It  is  also  an  interesting  fact,  as  bearing  on  sexual  selection,  that 
the  stridulating  organs  of  certain  male  Orthoptera  are  not  fully- 
developed  until  the  last  moult;  and  that  the  colors  of  certain 
male  dragon-flies  are  not  fully  developed  until  some  little  time 
after  their  emergence  from  the  pupal  state,  and  when  they  are 
ready  to  breed. 

Sexual  selection  implies  that  the  more  attractive  individuals 
are  preferred  by  the  opposite  sex;  and  as  with  insects,  when 
the  sexes  differ,  it  is  the  male  which,  with  some  rare  exceptions, 
is  the  more  ornamented,  and  departs  more  from  the  type  to 
which  the  species  belongs; — and  as  it  is  the  male  which  searches 
eagerly  for  the  female,  we  must  suppose  that  the  females  habitual- 
ly or  occasionally  prefer  the  more  beautiful  males,  and  that  these 
have  thus  acquired  their  beauty.  That  the  females  in  most  or 
all  the  orders  would  have  the  power  of  rejecting  any  particular 
male,  is  probable  from  the  many  singular  contrivances  possessed 
by  the  males,  such  as  great  jaws,  adhesive  cushions,  spines,  elon- 
gated legs,  &c.,  for  seizing  the  female;  for  these  contrivances 
show  that  there  is  some  diflaculty  in  the  act,  so  that  her  concur- 
rence would  seem  necessary.  Judging  from  what  we  know  of  the 
perceptive  powers  and  affections  of  various  insects,  there  is  no 
antecedent  improbability  in  sexual  selection  having  come  largely 
into  play;  but  we  have  as  yet  no  direct  evidence  on  this  head, 
and  some  facts  are  opposed  to  the  belief.  Nevertheless,  when 
we  see  many  males  pursuing  the  same  female,  we  can  hardly 
believe  that  the  pairing  is  left  to  blind  chance — that  the  female 
exerts  no  choice,  and  is  not  influenced  by  the  gorgeous  colors  or 
other  ornaments  with  which  the  male  is  decorated. 

If  we  admit  that  the  females  of  the  Homoptera  and  Orthoptera 
appreciate  the  musical  tones  of  their  male  partners,  and  that  the 
various  instruments  have  been  perfected  through  sexual  selec- 
tion, there  is  little  improbability  in  the  females  of  other  insects 
appreciating  beauty  in  form  or  color,  and  consequently  in  such 
characters  having  been  thus  gained  by  the  males.  But  from  the 
circumstance  of  color  being  so  variable,  and  from  its  having  been 
so  often  modified  for  the  sake  of  protection,  it  is  difficult  to  decide 
In  how  large  a  proportion  of  cases  sexual  selection  has  played 
a  part.  This  is  more  especially  difficult  in  those  Orders,  such  as 
Orthoptera,  Hymenoptera,  and  Coleoptera,  in  which  the  two 
sexes  rarely  differ  much  in  color;  for  we  are  then  left  to  mere 
analogy.  With  the  Coleoptera,  however,  as  before  remarked,  it 
is  in  the  great  Lamellicorn  group,  placed  by  some  authors  at 
the  head  of  the  Order,  and  in  which  we  sometimes  see  a  mutual 
attachment  between  the  sexes,  that  we  find  the  males  of  some 
species  possessing  weapons  for  sexual  strife,  others  furnished 
with  wonderful  horns,  many  with  stridulating  organs,  and  others 
ornamented  with  splendid  metallic  tints.    Hence  it  seems  prob- 

22 


326  THE  DESCENT  OF  MAN. 

able  that  all  these  characters  have  been  gained  through  the  same 
means,  namely,  sexual  selection.  With  butterflies  we  have  the 
best  evidence,  as  the  males  sometimes  take  pains  to  display  their 
beautiful  colors;  and  we  cannot  believe  that  they  would  act  thus, 
unless  the  display  was  of  use  to  them  in  their  courtship. 

When  we  treat  of  Birds,  we  shall  see  that  they  present  in 
their  secondary  sexual  characters  the  closest  analogy  with  in- 
sects. Thus,  many  male  birds  are  highly  pugnacious,  and  some 
are  furnished  with  special  weapons  for  fighting  with  their  rivals. 
They  possess  organs  which  are  used  during  the  breeding-season 
for  producing  vocal  and  instrumental  music.  They  are  frequently 
ornamented  with  combs,  horns,  wattles  and  plumes  of  the  most 
diversified  kinds,  and  are  decorated  with  beautiful  colors,  all 
evidently  for  the  sake  of  display.  We  shall  find  that,  as  with 
insects,  both  sexes  in  certain  groups  are  equally  beautiful  and 
are  equally  provided  with  ornaments  which  are  usually  confined 
to  the  male  sex.  In  other  groups  both  sexes  are  equally  plain- 
colored  and  unornamented.  Lastly,  in  some  few  anomalous  cases, 
the  females  are  more  beautiful  than  the  males.  We  shall  often 
find,  in  the  same  group  of  birds,  every  gradation  from  no  difference 
between  the  sexes,  to  an  extreme  difference.  We  shall  see  that 
female  birds,  like  female  insects,  often  possess  more  or  less  plain 
traces  or  rudiments  of  characters  which  properly  belong  to  the 
males  and  are  of  use  only  to  them.  The  analogy,  indeed,  in  all 
these  respects  between  birds  and  insects  is  curiously  close.  What- 
ever explanation  applies  to  the  one  class  probably  applies  to  the 
other;  and  this  explanation,  as  we  shall  hereafter  attempt  to 
show  in  further  detail,  is  sexual  selection. 


FISHES.  327 


CHAPTER  XII. 

SECONDARY  SEXUAL.  CHARACTERS  OF  FISHES,  AMPHIBIANS, 

AND    REPTILES. 

Fishes:  Courtship  and  battles  of  the  males— Larg-er  size  of  the  females 
— Males,  bright  colors  and  ornamental  appendages;  other  strange 
characters— Colors  and  appendages  acquired  by  the  males  during 
the  breeding-season  alone— Fishes  with  both  sexes  brilliantly  colored 
— Protective  colors— The  less  conspicuous  colors  of  the  female  can- 
not be  accounted  for  on  the  principle  of  protection — Male  fishes 
building  nests,  and  taking  charge  of  the  ova  and  young.  Amphib- 
ians: Differences  in  structure  and  color  between  the  sexes— Vocal 
organs.  Reptiles:  Chelonians— Crocodiles— Snakes,  colors  in  some 
cases  protective — Lizards,  battles  of — Ornamental  appendages- 
Strange  differences  in  structure  between  the  sexes— Colorsn-Sexual 
differences  almost  as  great  as  with  birds. 

We  have  now  arrived  at  the  great  sub-kingdom  of  the  Ver- 
tebrata,  and  will  commence  with  the  lowest  class,  that  of  Fishes. 
The  males  of  the  Plagiostomous  fishes  (sharks,  rays)  and  of 
ChimaBroid  fishes  are  provided  with  claspers  which  serve  to  retain 
the  female,  like  the  various  structures  possessed  by  many  of  the 
lower  animals.  Besides  the  claspers,  the  males  of  many  rays 
have  clusters  of  strong  sharp  spines  on  their  heads,  and  several 
rows  along  "the  upper  outer  surface  of  their  pectoral  fins."  These 
are  present  in  the  males  of  some  species,  which  have  other  parts 
of  their  bodies  smooth.  They  are  only  temporarily  developed 
during  the  breeding-season;  and  Dr.  Giinther  suspects  that  they 
are  brought  into  action  as  prehensile  organs  by  the  doubling  in- 
wards and  downwards  of  the  two  sides  of  the  body.  It  is  a  re- 
markable fact  that  the  females  and  not  the  males  of  some  spe- 
cies, as  of  Raia  clavata,  have  their  backs  studded  with  large  hook- 
formed  spines.^ 

The  males  alone  of  the  capelin  (Mallotus  villosus,  one  of  Sal- 
monidaB),  are  provided  with  a  ridge  of  closely-set,  brush-like 

lYarrell's  'Hist,  of  British  Fishes,'  vol.  ii.  1836,  pp.  417,  425,  436.  Dr. 
Gunther  informs  me  that  the  spines  in  R.  clavata  are  peculiar  to  the 
female, 


328  THE  DESCENT  OF  MAN. 

scales,  by  the  aid  of  which  two  males,  one  on  each  side,  hold  the 
female,  whilst  she  runs  with  great  swiftness  on  the  sandy  beach, 
and  there  deposits  her  spawn.^  The  widely  distinct  Monacanthus 
scopas  presents  a  somewhat  analogous  structure.  The  male,  as 
Dr.  Giinther  informs  me,  has  a  cluster  of  stiff,  straight  spines, 
like  those  of  a  comb,  on  the  sides  of  the  tail;  and  these  in  a 
specimen  six  inches  long  were  nearly  one  and  a  half  inches  in 
length;  the  female  has  in  the  same  place  a  cluster  of  bristles, 
which  may  be  compared  with  those  of  a  tooth-brush.  In  another 
species,  M.  peronii,  the  male  has  a  brush  like  that  possessed  by 
the  female  of  the  last  species,  whilst  the  sides  of  the  tail  in  the 
female  are  smooth.  In  some  other  species  of  the  same  genus  the 
tail  can  be  perceived  to  be  a  little  roughened  in  the  male  and  per- 
fectly smooth  in  the  female;  and  lastly  in  others,  both  sexes  have 
smooth  sides. 

The  males  of  many  fish  fight  for  the  possession  of  the  females. 
Thus  the  male  stickleback  (Gasterosteus  leiurus)  has  been  de- 
scribed as  "mad  with  delight,"  when  the  female  comes  out  of  her 
hiding-place  and  surveys  the  nest  which  he  has  made  for  her. 
"He  darts  round  her  in  every  direction,  then  to  his  accumulated 
"materials  for  the  nest,  then  back  again  in  an  instant;  and  as 
"she  does  not  advance  he  endeavors  to  push  her  with  his  snout, 
"and  then  tries  to  pull  her  by  the  tail  and  side  spine  to  the  nest.'" 

The  males  are  said  to  be  polygamists;*  they  are  extraordinarily 
bold  and  pugnacious,  whilst  "the  females  are  quite  pacific."  Their 
battles  are  at  times  desperate;  "for  these  puny  combatants  fasten 
"tight  on  each  other  for  several  seconds,  tumbling  over  and  over 
"again,  until  their  strength  appears  completely  exhausted."  With 
the  rough-tailed  stickleback  (G.  trachurus)  the  males  whilst  fight- 
ing swim  round  and  round  each  other,  biting  and  endeavoring  to 
pierce  each  other  with  their  raised  lateral  spines.  The  same 
writer  adds,^  "the  bite  of  these  little  furies  is  very  severe.  They 
"also  use  their  lateral  spines  with  such  fatal  effect,  that  I  have 
"seen  one  during  a  battle  absolutely  rip  his  opponent  quite  open, 
"so  that  he  sank  to  the  bottom  and  died."  When  a  fish  is  con- 
quered, "his  gallant  bearing  forsakes  him;  his  gay  colors  fade 
"away;  and  he  hides  his  disgrace  among  his  peaceable  compan- 
"ions,  but  is  for  some  time  the  constant  object  of  his  conqueror's 
"persecution." 

The  male  salmon  is  as  pugnacious  as  the  little  stickleback; 
and  so  is  the  male  trout,  as  I  hear  from  Dr.  Giinther.    Mr.  Shaw 


^  'The  American  Naturalist,'  April,  1871,  p.  119. 

3  See  Mr.  R.  Warington's  interesting  articles  in  'Annals  and  Mag-,  of 
Nat.  Hist.'  Oct.  1S52,  and  Nov.,  1855. 
*  Noel  Humphreys,  'River  Gardens,'  1857. 
5  Loudon's  'Mag.  of  Nat.  History,'  vol.  iii.  1830,  p.  331. 


FISHES. 


32y 


saw  a  violent  contest  between  two  male  salmon  which  lasted  the 
whole  day;  and  Mr.  R.  Buist,  Superintendent  of  Fisheries,  in- 
forms me  that  he  has  often  watched  from  the  bridge  at  Perth  the 
males  driving  away  their  rivals,  whilst  the  females  were  spawn- 


Fig.  27.    Head  of  male  common  salmon  (Salmo  salar)  during  the 

breeding-season. 

[This  drawing,  as  well  as  all  the  others  in  the  present  chapter,  have 
been  executed  by  the  well-known  artist,  Mr.  G.  Ford,  from  specimens 
in  the  British  Museum,  under  the  kind  superintendence  of  Dr.  Gun- 
ther.] 

ing.  The  males  "are  constantly  fighting  and  tearing  each  other 
"on  the  spawning-beds,  and  many  so  injure  each  other  as  to 
"cause  the  death  of  numbers,  many  being  seen  swimming  near 
"the  banks  of  the  river  in  a  state  of  exhaustion,  and  apparently 


330  THE  DESCENT  OP'  MAN. 

*'iii  a  dying  state."®  Mr.  Buist  informs  me,  that  in  June,  1868, 
the  keeper  of  the  Stormontfield  breeding-ponds  visited  the  north- 
ern Tyne  and  found  about  300  dead  salmon,  all  of  which  with  one 
exception  were  males;  and  he  was  convinced  that  they  had  lost 
their  lives  by  fighting. 


Fig".  28.     Head  of  female  salmon. 

The  most  curious  point  about  the  male  sahr^on  is  that  during 
the  breeding-season,  besides  a  slight  change  in  color,  "the  lower 
"jaw  elongates,  and  a  cartilaginous  projection  turns  upwards 
"from  the  point,  which,  when  the  jaws  are  closed,  occupies  a 
"deep  cavity  between  the  intermaxillary  bones  of  the  upper  jaw."^ 
(Figs.  27  and  28.)     In  our  salmon  this  change  of  structure  lasts 

«  'The  Field,'  June  29th,  1867.  For  Mr.  Shaw's  statement,  see  'Edin- 
burgti  Review,'  1843.  Another  experienced  observer  (Scrope's  'Days  of 
Salmon  Fisihing-,'  p.  60)  remarks  that  like  the  stag,  the  male  would,' if 
he  could,  keep  all  other  males  away. 

"^  Yarrell,  'History  of  British  Fishes,'  vol.  ii.  1836,  p.  10. 


FISHES.  331 

only  during  the  breeding-season;  but  in  the  Salmo  lycaodon  of 
N.-W.  America  the  change,  as  Mr.  J.  K.  Lord**  believes,  is  perma- 
nent, and  best  marked  in  the  older  males  which  have  previously 
ascended  the  rivers.  In  these  old  males  the  jaw  becomes  de- 
veloped into  an  immense  hook-like  projection,  and  the  teeth 
grow  into  regular  fangs,  often  more  than  half  an  inch  in  length. 
With  the  European  salmon,  according  to  Mr.  Lloyd,^  the  tem- 
porary hook-like  structure  serves  to  strengthen  and  protect  the 
jaws,  when  one  male  charges  another  with  wonderful  violence; 
but  the  greatly  developed  teeth  of  the  male  American  salmon 
may  be  compared  v/ith  the  tusks  of  many  male  mammals,  and 
they  indicate  an  offensive  rather  than  a  protective  purpose. 

The  salmon  is  not  the  only  fish  in  which. the  teeth  differ  in 
the  two  sexes;  as  this  is  the  case  with  many  rays.  In  the 
thornback  (Raia  clavata)  the  adult  male  has  sharp,  pointed 
teeth,  directed  backwards,  whilst  those  of  the  female  are  broad 
and  flat,  and  form  a  pavement;  so  that  these  teeth  differ  in  the 
two  sexes  of  the  same  species  more  than  is  usual  in  distinct 
genera  of  the  same  family.  The  teeth  of  the  male  become  sharp 
only  when  he  is  adult;  whilst  young  they  are  broad  and  flat 
like  those  of  the  female.  As  so  frequently  occurs  with  secondary 
sexual  characters,  both  sexes  of  seme  species  of  rays  (for  instance 
R.  batis),  when  adult,  possess  sharp  pointed  teeth;  and  here  a 
character,  proper  to  and  primarily  gained  by  the  male,  appears 
to  have  been  transmitted  to  the  offspring  of  both  sexes.  The 
teeth  are  likewise  pointed  in  both  sexes  of  R.  maculata,  but  only 
when  quite  adult;  the  males  acquiring  them  at  an  earlier  age 
than  the  females.  We  shall  hereafter  meet  with  analogous  cases 
in  certain  birds,  in  which  the  male  acquires  the  plumage  common 
to  both  sexes  when  adult,  at  a  somewhat  earlier  age  than  does 
the  female.  With  other  species  of  rays  the  males  even  when  old 
never  possess  sharp  teeth,  and  consequently  the  adults  of  both 
sexes  are  provided  with  broad,  flat  teeth  like  those  of  the  young, 
and  like  those  of  the  mature  females  of  the  above-mentionea 
species.^"  As  the  rays  are  bold,  strong  and  voracious  fish,  we  may 
suspect  that  the  males  require  their  sharp  teeth  for  fighting  with 
their  rivals;  but  as  they  possess  many  parts  modified  and  adapted 
for  the  prehension  of  the  female,  it  is  possible  that  their  teeth 
may  be  used  for  this  purpose. 

In  regard  to  size,  M.  Carbonnier"  maintains  that  the  female  of 
almost  all  fishes  is  larger  than  the  male;    and  Dr.  Giinther  does 

8  'The  Naturalist  in  Vancouver's  Island,'  vol.  i.  1866,  p.  54. 

9  'Scandinavian  Adventures,'  vol.  i.  1854,  pp.  100,  104. 

10  See  Yarrell's  account  of  the  rays  in  his  'Hist,  of  British  Fisihe®,* 
vol.  ii.  1836,  p.  416,  with  an  excellent  figure,  and  p.  422,  432. 

"  As  quoted  in  'The  Farmer,'  1868,  p.  369. 


332  THE  DESCENT  OF  MAN. 

not  know  of  a  single  instance  in  which  the  aale  is  actually 
larger  than  the  female.  With  some  Cyprinodonts  the  male  is 
not  even  half  as  large.  As  in  many  kinds  of  fishes  the  males 
habitually  fight  together,  it  is  surprising  that  they  have  not  gen- 
erally become  larger  and  stronger  than  the  females  through  the 
effects  of  sexual  selection.  The  males  suffer  from  their  small 
size,  for  according  to  M.  Carbonnier,  they  are  liable  to  be  de- 
voured by  the  females  of  their  own  species  when  carnivorous. 


Fig.  29.    Callionymus  lyra.    Upper  figure,  male;    lower  figure,  female. 
N.  B.     The  lower  figure  is  more  reduced  than  the  upper. 

and  no  doubt  by  other  species^  Increased  size  must  be  in  some 
manner  of  more  importance  to  the  females,  than  strength  and 
size  are  to  the  males  for  fighting  with  other  males;  and  this 
perhaps  is  to  allow  of  the  production  of  a  vast  number  of  ova. 

In  many  species  the  male  alone  is  ornamented  with  bright 
colors;  or  these  are  much  brighter  in  the  male  than  the  female. 
The  male,  also,  is  sometimes  provided  with  appendages  which  ap- 
pear to  be  of  no  more  use  to  him  for  the  ordinary  purposes  of 
life,  than  are  the  tail  feathers  to  the  peacock.  I  am  indebted 
for  most  of  the  following  facts  to  the  kindness  of  Dr.  Gunther. 
There  is  reason  to  suspect  that  many  tropical  fishes  differ  sexually 
in  color  and  structure;  and  there  are  some  striking  cases  with 
our  British  fishes.    The  male  Callionymus  lyra  has  been  called 


FISHES. 


333 


the  gemmeous  dragonet  "from  its  brilliant  gem-like  colors." 
When  fresh  caught  from  the  sea  the  body  is  yellow  of  various 
shades,  striped  and  spotted  with  vivid  blue  on  the  head;  the 
dorsal  fins  are  pale  brown  with  dark  longitudinal  bands;  the 
ventral,  caudal,  and  anal  fins  being  bluish-black.  The  female, 
or  sordid  dragonet,  was  considered  by  Linnasus,  and  by  many 
subsequent  naturalists,  as  a  distinct  species;  it  is  of  a  dingy  red- 
dish-brown, with  the  dorsal  fin  brown  and  the  other  fins  white. 


Fig. 


30.    Xiphophorus  Hellerii.    Upper  figure,  male; 

female. 


lower  figure. 


The  sexes  differ  also  in  the  proportional  size  of  the  head  and 
mouth,  and  in  the  position  of  the  eyes;^^  i^^i  i\^q  most  striking  dif- 
ference is  the  extraordinary  elongation  in  the  male  (fig.  29)  of 
the  dorsal  fin.  Mr.  W.  Saville  Kent  remarks  that  this  "singular 
"appendage  appears  from  my  observations  of  the  species  in  con- 
"finement,  to  be  subservient  to  the  same  end  as  the  wattles, 
"crests,  and  other  abnormal  adjuncts  of  the  male  in  gallinaceous 
"birds,  for  the  purpose  of  fascinating  their  mates.""  The  young 
males  resemble  the  adult  females  in  structure  and  color.  Through- 
out the  genus  Callionymus,"  the  male  is  generally  much  more 
brightly  spotted  than  the  female,  and  in  several  species,  not  only 
the  dorsal,  but  the  anal  fin  is  much  elongated  in  the  males. 


12  1  have  drawn  up  this  description  from  Tarrell's  'British  Fishes,' 
vol.  i.  1836,  pp.  261  and  266. 

13  'Nature,'  July,  1873,  p.  264. 

"  'Catalogue  of  Acanth.    Fishes  in  the  British  Museum,'  by  Dr.  Gun- 
ther,  1861,  pp.  138-151. 


334  THE  DESCENT  OF  MAN. 

The  male  of  the  Cottus  scorpius,  or  sea-scorpion,  is  slenderer 
and  smaller  than  the  female.  There  is  also  a  great  difference 
in  color  between  them.  It  is  difficult,  as  Mr.  Lloyd^^  remarks, 
"for  any  one,  who  has  not  seen  this  fish  during  the  spawning- 
"season,  when  its  hues  are  brightest,  to  conceive  the  admixture 
"of  brilliant  colors  with  which  it,  in  other  respects  so  ill-favored, 
**is  at  that  time  adorned."  Both  sexes  of  the  Labrus  mixtus, 
although  very  different  in  color,  are  beautiful;  the  male  being 
orange  with  bright  blue  stripes,  and  the  female  bright  red  with 
some  black  spots  on  the  back. 

In  the  very  distinct  family  of  the  Cyprinodontid® — inhabitants 
of  the  fresh  waters  of  foreign  lands — the  sexes  sometimes  differ 
much  in  various  characters.  In  the  male  of  the  Mollienesia  peten- 
ensis,^®  the  dorsal  fin  is  greatly  developed  and  is  marked  with  a 
row  of  large,  round,  ocellated,  bright-colored  spots;  whilst  the 
same  fin  in  the  female  is  smaller,  of  a  different  shape,  and 
marked  only  with  irregularly  curved  brown  spots.  In  the  male 
the  basal  margin  of  the  anal  fin  is  also  a  little  produced  and  dark 
colored.  In  the  male  of  an  allied  form,  the  Xiphophorus  Hellerii 
(fig.  30),  the  inferior  margin  of  the  caudal  fin  is  developed  into 
a  long  filament,  which,  as  I  hear  from  Dr.  Giinther,  is  striped  with 
bright  colors.  This  filament  does  not  contain  any  muscles,  and 
apparently  cannot  be  of  any  direct  use  to  the  fish.  As  in  the 
case  of  the  Callionymus,  the  males  whilst  young  resemble  the 
adult  females  in  color  and  structure.  Sexual  differences  such  as 
these  may  be  strictly  compared  with  those  which  are  so  frequent 
with  gallinaceous  birds." 

In  a  siluroid  fish,  inhabiting  the  fresh  waters  of  South  America, 
the  Plecostomus  barbatus^^  (fig.  31),  the  male  has  its  mouth  and 
inter- operculum  fringed  with  a  beard  of  stiff  hairs,  of  which  the 
female  shows  hardly  a  trace.  These  hairs  are  of  the  nature  of 
scales.  In  another  species  of  the  same  genus,  soft  flexible  ten- 
tacles project  from  the  front  part  of  the  head  of  the  male,  which 
are  absent  in  the  female.  These  tentacles  are  prolongations  of 
the  true  skin,  and  therefore  are  not  homologous  with  the  stiff 
hairs  of  the  former  species;  but  it  can  hardly  be  doubted  that 
both  serve  the  same  purpose.  What  this  purpose  may  be,  it  is 
difficult  to  conjecture;  ornament  does  not  here  seem  probable, 
but  we  can  hardly  suppose  that  stiff  hairs  and  flexible  filaments 
can  be  useful  in  any  ordinary  way  to  the  males  alone.    In  that 

15  'Game  Birds  of  Sweden,'  &c.,  1867,  p.  466. 

10  With  respect  to  this  and  the  following-  species  I  am  indebted  to 
Dr.  Gunther  for  information:  see,  also,  his  paper  on  the  'Fishes  of 
Central  America,'  in  'Transact.  Zoolog-.  Soc'  vol.  vi.  1868,  p.  485. 

"  Dr.  Gunther  makes  this  remark;  'Catalogue  of  Fishes  in  the  Britisli 
Museum,'  vol.  iii.  1861,  p.  141. 

""3  See  Dr.  Gunther  on  this  genus,  in  'Proc.  Zoolog.  Soc'  1868,  p.  232. 


FISHES. 


335 


Fig-.   31.     Plecostomus  barbatus.    Upper  figure,  head  of  male;   lower 

figure,  female. 


336  THE  DESCENT  OF  MAN. 

strange  monster,  the  Chimaera  monstrosa,  the  male  has  a  hook- 
shaped  bone  on  the  top  of  the  head,  directed  forwards,  with  its 
end  rounded  and  covered  with  sharp  spines;  in  the  female  "this 
"crown  is  altogether  absent,"  but  what  its  use  may  be  to  the  male 
is  utterly  unknown.^® 

The  structures  as  yet  referred  to  are  permanent  in  the  male 
after  he  has  arrived  at  maturity;  but  with  some  Blennies,  and  in 
another  allied  genus,^''  a  crest  is  developed  on  the  head  of  the 
male  only  during  the  breeding-season,  and  the  body  at  the  same 
time  becomes  more  brightly-colored.  There  can  be  little  doubt 
that  this  crest  serves  as  a  temporary  sexual  ornament,  for  the 
female  does  not  exhibit  a  trace  of  it.  In  other  species  of  the 
same  genus  both  sexes  possess  a  crest,  and  in  at  least  one  species 
neither  sex  is  thus  provided.  In  many  of  the  Chromidse,  for 
instance  in  Geophagus  and  especially  in  Cichla,  the  males,  as  I 
hear  from  Professor  Agassiz,^^  have  a  conspicuous  protuberance 
on  the  forehead,  which  is  wholly  wanting  in  the  females  and  in 
the  young  males.  Professor  Agassiz  adds,  "I  have  often  observed 
"these  fishes  at  the  time  of  spawning  when  the  protuberance  is 
"largest,  and  at  other  seasons  when  it  is  totally  wanting,  and 
"the  two  sexes  show  no  difference  whatever  in  the  outline  of 
"the  profile  of  the  head.  I  never  could  ascertain  that  it  subserves 
"any  special  function,  and  the  Indians  on  the  Amazon  know 
"nothing  about  its  use."  These  protuberances  resemble,  in  their 
periodical  appearance,  the  fleshy  caruncles  on  the  heads  of  certain 
birds;  but  whether  they  serve  as  ornaments  must  remain  at 
present  doubtful. 

I  hear  from  Professor  Agassiz  and  Dr.  Giinther,  that  the  males 
of  those  fishes,  which  differ  permanently  in  color  from  the  fe- 
males, often  become  more  brilliant  during  the  breeding-season. 
This  is  likewise  the  case  with  a  multitude  of  fishes,  the  sexes  of 
which  are  identical  in  color  at  all  other  seasons  of  the  year.  The 
tench,  roach,  and  perch  may  be  given  as  instances.  The  male 
salmon  at  this  season  is  "marked  on  the  cheeks  with  orange- 
"colored  stripes,  which  give  it  the  appearance  of  a  Labrus,  and 
"the  body  partakes  of  a  golden  orange  tinge.  The  females  are 
"dark  in  color,  and  are  commonly  called  black-fish."^-  An  anal- 
ogous and  even  greater  change  takes  place  with  the  Salmo  eriox 
or  bull  trout;    the  males  of  the  char  (S.  umbla)  are  likewise  at 

i»  F.  Buckland,  in  'Land  and  Water,'  July,  1868,  p.  377,  with  a  figure. 
Many  other  cases  could  be  added  of  structures  peculiar  to  the  male, 
of  which  the  uses  are  not  known. 

20  Dr.  Gunther,  'Catalogue  of  Fishes,'  vol.  iii.  pp.  221  and  240. 

21  See,  also,  'A  Journey  in  Brazil,'  by  Prof,  and  Mrs.  Agassiz,  1868, 
p.  220. 

M  Yarrell,  'British  Fishes,'  vol.  i  1836,  pp.  10,  12,  35. 


FISHES.  337 

this  season  rather  brighter  in  color  than  the  females.-^  The 
colors  of  the  pike  (Esox  reticulatus)  of  the  United  States,  especi- 
ally of  the  male,  become,  during  the  breeding-season,  exceedingly- 
intense,  brilliant,  and  iridescent.-*  Another  striking  instance  out 
of  many  is  afforded  by  the  male  stickleback  (Gasterosteus  leiu- 
rus),  which  is  described  by  Mr.  Warington,-^  as  being  then  "beau- 
"tifUi  beyond  description."  The  back  and  eyes  of  the  female  are 
simply  brown,  and  the  belly  white.  The  eyes  of  the  male,  on 
the  other  hand,  are  "of  the  most  splendid  green,  having  a  metallic 
"luster  like  the  green  feathers  of  some  humming-birds.  The 
"throat  and  belly  are  of  a  bright  crimson,  the  back  of  an  ashy- 
"green,  and  the  whole  fish  appears  as  though  it  were  somewhat 
"translucent  and  glowed  with  an  internal  incandescence."  After 
the  breeding-season  these  colors  all  change,  the  throat  and  belly 
become  of  a  paler  red,  the  back  more  green,  and  the  glowing  tints 
subside. 

With  respect  to  the  courtship  of  fishes,  other  cases  have  been 
observed  since  the  first  edition  of  this  book  appeared,  besides  that 
already  given  of  the  stickleback.  Mr.  W.  S.  Kent  says  that  the 
male  of  the  Labrus  mixtus,  which,  as  we  have  seen,  differs  in 
color  from  the  female,  makes  "a  deep  hollow  in  the  sand  of  the 
"tank,  and  then  endeavors  in  the  most  persuasive  manner  to  in- 
"duce  a  female  of  the  same  species  to  share  it  with  him,  swim- 
"ming  backwards  and  forwards  between  her  and  the  completed 
"nest,  and  plainly  exhibiting  the  greatest  anxiety  for  her  to  fol- 
"low."  The  males  of  the  Cantharus  lineatus  become,  during  the 
breeding-season,  of  deep  leaden-black;  they  then  retire  from 
the  shoal,  and  excavate  a  hollow  as  a  nest.  "Bach  male  now 
"mounts  vigilant  guard  over  his  respective  hollow,  and  vigor- 
"ously  attacks  and  drives  away  any  other  fish  of  the  same  sex. 
"Towards  his  companions  of  the  opposite  sex  his  conduct  is  far 
"different;  many  of  the  latter  are  now  distended  with  spawn,  and 
"these  he  endeavors  by  all  the  means  in  his  power  to  lure  singly 
"to  his  prepared  hollow,  and  there  to  deposit  the  myriad  ova  witb 
"which  they  are  laden,  which  he  then  protects  and  guards  with 
"the  greatest  care."^" 

A  more  striking  case  of  courtship,  as  well  as  of  display,  by  the 
males  of  a  Chinese  Macropus  has  been  given  by  M,  Carbonnier, 
who  carefully  observed  these  fishes  under  confinement."  The 
males  are  most  beautifully  colored,  more  so  than  the  females. 
During  the  breeding-season  they  contend  for  the  possession  of 

23  W.  Thompson,  in  'Annals  and  Mag.  of  Nat.  History,'  vol.  vi.  1841, 
p.  440. 
"*  'The  American  Agriculturist,'  1868,  p.  100. 
23  'Annals  and  Mag.  of  Nat.  Hist.'  Oct.  1852. 
2«  'Nature,'  May,  1873,  p.  25. 

27  'Bull,  de  la  Soc.  d'Acclimat.'  Paris,  July,  1869,  and  Jan.  1870. 
23 


338  THE  DESCENT  OF  MAN. 

the  females,  and,  in  the  act  of  courtship,  expand  their  fins, 
which  are  spotted  and  ornamented  with  brightly  colored  rays, 
in  the  same  manner,  according  to  M.  Carbonnier,  as  the  peacock. 
They  then  also  bound  about  the  females  with  much  vivacity,  and 
appear  by  "I'etalage  de  leurs  vives  couleurs  chercher  a  attirer 
"I'attention  des  femelles,  lesquelles  ne  paraissaient  indifferentes 
"a  ce  manege,  elles  nageaient  avec  une  molle  lenteur  vers  les 
"males  et  semblaient  se  complaire  dans  leur  voisinage."  After 
the  male  has  won  his  bride,  he  makes  a  little  disc  of  froth  by 
blowing  air  and  mucus  out  of  his  mouth.  He  then  collects  the 
fertilized  ova,  dropped  by  the  female,  in  his  mouth;  and  this 
caused  M.  Carbonnier  much  alarm,  as  he  thought  that  they  were 
going  to  be  devoured.  But  the  male  soon  deposits  them  in  the 
disc  of  froth,  afterwards  guarding  them,  repairing  the  froth,  and 
taking  care  of  the  young  when  hatched.  I  mention  these  par- 
ticulars because,  as  *we  shall  presently  see,  there  are  fishes,  the 
males  of  which  hatch  their  eggs  in  their  mouths;  and  those  who 
do  not  believe  in  the  principle  of  gradual  evolution  might  ask 
how  could  such  a  habit  have  originated;  but  the  difQculty  is 
much  diminished  when  we  know  that  there  are  fishes  which  thus 
collect  and  carry  the  eggs;  for  if  delayed  by  any  cause  in  de- 
positing them,  the  habit  of  hatching  them  in  their  mouths  might 
have  been  acquired. 

To  return  to  our  more  immediate  subject.  The  case  stands 
thus:  female  fishes,  as  far  as  I  can  learn,  never  willingly  spawn 
except  in  the  presence  of  the  males;  and  the  males  never  fertilize 
the  ova  except  in  the  presence  of  the  females.  The  males  fight 
for  the  possession  of  the  females.  In  many  species,  the  males 
whilst  young  resemble  the  females  in  color;  but  when  adult 
become  much  more  brilliant,  and  retain  their  colors  throughout 
life.  In  other  species  the  males  become  brighter  than  the  females 
and  otherwise  more  highly  ornamented,  only  during  the  season 
of  love.  The  males  sedulously  court  the  females,  and  in  one 
case,  as  we  have  seen,  take  pains  in  displaying  their  beauty 
before  them.  Can  it  be  believed  that  they  would  thus  act  to  no 
purpose  during  their  courtship?  And  this  would  be  the  case, 
unless  the  females  exert  some  choice  and  select  those  males  which 
please  or  excite  them  most.  If  the  female  exerts  such  choice,  all 
the  above  facts  on  the  ornamentation  of  the  males  become  at  once 
intelligible  by  the  aid  of  sexual  selection. 

We  have  next  to  inquire  whether  this  view  of  the  bright  colors 
of  certain  male  fishes  having  been  acquired  through  sexual  selec- 
tion can,  through  the  law  of  the  equal  transmission  of  characters 
to  both  sexes,  be  extended  to  those  groups  in  which  the  males  and 
females  are  brilliant  in  the  same,  or  nearly  the  same  degree  and 
manner.  In  such  a  genus  as  Labrus,  which  includes  some  of  the 
most  splendid   fishes   in  the  world — for  instance,   the   Peacock 


FISHES.  339 

Labrus  (L.  pavo),  described,^*  with  pardonable  exaggeration,  as 
formed  of  polished  scales  of  gold,  encrusting  lapis-lazuli,  rubies, 
sapphires,  emeralds,  and  amethysts — we  may,  with  much  prob- 
ability, accept  this  belief;  for  we  have  seen  that  the  sexes  in 
at  least  one  species  of  the  genus  differ  greatly  in  color.  With  some 
fishes,  as  with  many  of  the  lowest  animals,  splendid  colors  may 
be  the  direct  result  of  the  nature  of  their  tissues  and  of  the  sur- 
rounding conditions,  without  the  aid  of  selection  of  any  kind. 
The  gold-fish  (Cyprinus  auratus),  judging  from  the  analogy  of  the 
golden  variety  of  the  common  carp,  is  perhaps  a  case  in  point, 
as  it  may  owe  its  splendid  colors  to  a  single  abrupt  variation, 
due  to  the  conditions  to  which  this  fish  has  been  subjected  under 
confinement.  It  is,  however,  more  probable  that  these  colors 
have  been  intensified  through  artificial  selection,  as  this  species 
has  been  carefully  bred  in  China  from  a  remote  period.^*  Under 
natural  conditions  it  does  not  seem  probable  that  beings  so  highly 
organized  as  fishes,  and  which  live  under  such  complex  rela- 
tions, should  become  brilliantly  colored  without  suffering  some 
evil  or  receiving  some  benefit  from  so  great  a  change,  and  con- 
sequently without  the  intervention  of  natural  selection. 

What,  then,  are  we  to  conclude  in  regard  to  the  many  fishes, 
both  sexes  of  which  are  splendidly  colored?  Mr.  Wallace'"  be- 
lieves that  the  species  which  frequent  reefs,  where  corals  and 
other  brightly-colored  organisms  abound,  are  brightly  colored  in 
order  to  escape  detection  by  their  enemies;  but  according  to  my 
recollection  they  were  thus  rendered  highly  conspicuous.  In 
the  fresh-waters  of  the  tropics  there  are  no  brilliantly-colored 
corals  or  other  organisms  for  the  fishes  to  resemble;  yet  many 
species  in  the  Amazons  are  beautifully  colored,  and  many  of  the 
carnivorous  Cyprinidae  in  India  are  ornamented  with  "bright 
"longitudinal  lines  of  various  tints."^^  Mr.  M'Clelland,  in  de- 
scribing these  fishes,  goes  so  far  as  to  suppose  that  "the  peculiar 

28  Bory  de  Saint  Vincent,  in  'Diet.  Class.  d'Hist.  Nat.'  torn.  ix.  1826, 
p.  151. 

-9  Owing  to  some  remarks  on  this  subject,  made  in  my  work  'On  the 
Variation  of  Animals  under  Domestication,'  Mr.  W.  F.  Mayers  ('Chi- 
nese Notes  and  Queries,'  Aug.  1868,  p.  123)  has  searched  the  ancient 
Chinese  encyclopedias.  He  finds  that  gold-fish  were  first  reared  in  con- 
finement during  the  Sung  Dynasty,  which  commenced  A.  D.  960.  In 
the  year  1129  these  fishes  abounded.  In  another  place  it  is  said  that 
since  the  year  1548  there  has  been  "produced  at  Hangchow  a  variety 
"called  the  fire-fish,  from  its  intensely  red  color.  It  is  universally  ad- 
"mired,  and  there  is  not  a  household  where  it  is  not  cultivated,  in 
"rivalry  as  to  its  color,  and  as  a  source  of  profit." 

30  'Westminster  Review,'  July,  1867,  p.  7. 

81  'Indian  Cyprinidae,'  by  Mr.  J.  M'Clelland,  'Asiatic  Researches,'  vol. 
Xts.  part  ii.  1839,  p.  230. 


340  THE  DESCENT  OF  MAN. 

"brilliancy  of  their  colors"  serves  as  "a  better  mark  for  king- 
"fishers,  terns,  and  other  birds  which  are  destined  to  keep  the 
"number  of  these  fishes  in  check;"  but  at  the  present  day  few 
naturalists  will  admit  that  any  animal  has  been  made  conspicuous 
as  an  aid  to  its  own  destruction.  It  is  possible  that  certain  fishes 
may  have  been  rendered  conspicuous  in  order  to  warn  birds  and 
beasts  of  prey  that  they  were  unpalatable,  as  explained  when 
treating  of  caterpillars;  but  it  is  not,  I  believe,  known  that  any 
fish,  at  least  any  fresh-water  fish,  is  rejected  from  being  distaste- 
ful to  fish-devouring  animals.  On  the  whole,  the  most  probable 
view  in  regard  to  the  fishes,  of  which  both  sexes  are  brilliantly 
colored,  is  that  their  colors  were  acquired  by  the  males  as  a 
sexual  ornament,  and  were  transferred  equally,  or  nearly  so,  to 
the  other  sex. 

We  have  now  to  consider  whether,  when  the  male  differs  in  a 
marked  manner  from  the  female  in  color  or  in  other  ornaments, 
he  alone  has  been  modified,  the  variations  being  inherited  by  his 
male  offspring  alone;  or  whether  the  female  has  been  specially 
modified  and  rendered  inconspicuous  for  the  sake  of  protection, 
such  modifications  being  inherited  only  by  the  females.  It  is 
impossible  to  doubt  that  color  has  been  gained  by  many  fishes 
as  a  protection:  no  one  can  examine  the  speckled  upper  surface 
of  a  flounder,  and  overlook  its  resemblance  to  the  sandy  bed  of 
the  sea  on  which  it  lives.  Certain  fishes,  moreover,  can  through 
the  action  of  the  nervous  system,  change  their  colors  in  adapta- 
tion to  surrounding  objects,  and  that  within  a  short  time.^-  One 
of  the  most  striking  instances  ever  recorded  of  an  animal  being 
protected  by  its  color  (as  far  as  it  can  be  judged  of  in  preserved 
specimens),  as  well  as  by  its  form,  is  that  given  by  Dr.  Giinther^^* 
of  a  pipe-fish,  which,  with  its  reddish  streaming  filanients,  is 
hardly  distinguishable  from  the  sea- weed  to  which  it  clings  with 
its  prehensile  tail.  But  the  question  now  under  consideration  is 
whether  the  females  alone  have  been  modified  for  this  object. 
We  can  see  that  one  sex  will  not  be  modified  through  natural  se- 
lection for  the  sake  of  protection  more  than  the  other,  supposing 
both  to  vary,  unless  one  sex  is  exposed  for  a  longer  period  to 
danger,  or  has  less  power  of  escaping  from  such  danger  than  the 
other;  and  it  does  not  appear  that  with  fishes  the  sexes  differ  in 
these  respects.  As  far  as  there  is  any  difference,  the  males,  from 
being  generally  smaller  and  from  wandering  more  about,  are 
exposed  to  greater  danger  than  the  females;  and  yet,  when  the 
sexes  differ,  the  males  are  almost  always  the  more  conspicuously 
colored.  The  ova  are  fertilized  immediately  after  being  deposited; 
and  when  this  process  lasts  for  several  days,  as  in  the  case  of 

82  G.  Pouchet,  L'Institut,  Nov.  1,  1871,  p.  134. 

83  'Proc.  Zoolog.  Soc'  1865,  p.  327,  pi.  xiv.  and  xv. 


FISHES.  341 

the  salmon/*  the  female,  during  the  whole  time,  is  attended  by 
the  male.  After  the  ova  are  fertilized  they  are,  in  most  cases, 
left  unprotected  by  both  parents,  so  that  the  males  and  females, 
as  far  as  oviposition  is  concerned,  are  equally  exposed  to  danger, 
and  both  are  equally  important  for  the  production  of  fertile  ova; 
consequently  the  more  or  less  brightly-colored  individuals  of 
either  sex  would  be  equally  liable  to  be  destroyed  or  preserved, 
and  both  would  have  an  equal  influence  on  the  colors  of  their  off- 
spring. 

Certain  fishes,  belonging  to  several  families,  make  nests,  and 
some  of  them  take  care  of  their  young  when  hatched.  Both 
sexes  of  the  bright  colored  Crenilabrus  massa  and  melops  work 
together  in  building  their  nests  with  sea-weed,  shells,  &c.^^  But 
the  males  of  certain  fishes  do  all  the  work,  and  afterwards  take 
exclusive  charge  of  the  young.  This  is  the  case  with  the  dull- 
colored  gobies,"*'  in  which  the  sexes  are  not  known  to  differ  in 
color,  and  likewise  with  the  sticklebacks  (Gasterosteus),  in 
which  the  males  become  brilliantly  colored  during  the  spawning 
season.  The  male  of  the  smooth-tailed  stickleback  (G.  leiurus) 
performs  the  duties  of  a  nurse  with  exemplary  care  and  vigilance 
during  a  long  time,  and  is  continually  employed  in  gently  leading 
back  the  young  to  the  nest,  when  they  stray  too  far.  He  courage- 
ously drives  away  all  enemies,  including  the  females  of  his  own 
species.  It  would  indeed  be  no  small  relief  to  the  male,  if  the  fe- 
male, after  depositing  her  eggs,  were  immediately  devoured  by 
some  enemy,  for  he  is  forced  incessantly  to  drive  her  from  the 
nest." 

The  males  of  certain  other  fishes  inhabiting  South  America 
and  Ceylon,  belonging  to  two  distinct  Orders,  have  the  extraor- 
dinary habit  of  hatching  within  their  mouths  or  branchial  cavi- 
ties, the  eggs  laid  by  the  females.^^  I  am  informed  by  Professor 
Agassiz  that  the  males  of  the  Amazonian  species  which  follow  this 
habit,  "not  only  are  generally  brighter  than  the  females,  but 
"the  difference  is  greater  at  the  spawning-season  than  at  any 
"other  time."  The  species  of  Geophagus  act  in  the  same  manner; 
and  in  this  genus,  a  conspicuous  protuberance  becomes  developed 
on  the  forehead  of  the  males  during  the  breeding-season.    Witn 


3*  Yarrell,  'British  Fishes,'  vol.  ii.  p.  U. 

35  According-  to  the  observations  of  M.  Gerbe;  see  Gunther's  'Record 
of  Zoolog-.  Literature,'  1865,  p.  194. 

36  Cuvier,  'Regne  Animal,'  vol.  ii.  1829,  p.  242. 

37  See  Mr.  Warington' si  most  interesting-  description  of  the  habits  of 
the  Gasterosteus  leiurus,  in  'Annals  and  Mag-,  of  Nat.  Hist.'  Novem- 
ber, 1855. 

38  Prof.  Wyman,  in  'Proc.  Boston  Soc.  of  Nat.  Hist.'  Sept.  15,  1857. 
Also  Prof.  Turner,  in  'Journal  of  Anatomy  and  Phys.'  Nov.  1,  1866, 
p.  78.     Dr.  Gunther  has  likewise  described  other  cases. 

23 


342  THE  DESCENT  OF  MAN. 

the  various  species  of  Chromids,  as  Professor  Agassiz  likewise 
informs  me,  sexual  differences  in  color  may  be  observed,  "whether 
"they  lay  their  eggs  in  the  water  among  aquatic  plants,  or  deposit 
"them  in  holes,  leaving  them  to  come  out  without  further  care, 
"or  build  shallow  nests  in  the  river  mud,  over  which  they  sit,  as 
"our  Pomotis  does.  It  ought  also  to  be  observed  that  these  sitters 
"are  among  the  brightest  s-pecies  in  their  respective  families;  for 
"instance,  Hygrogonus  is  bright  green,  with  large  black  ocelli, 
"encircled  with  the  most  brilliant  red."  Whether  with  all  the 
species  of  Chromids  it  is  the  male  alone  which  sits  on  the  eggs  is 
not  known.  It  is,  however,  manifest  that  the  fact  of  the  eggs 
being  protected  or  unprotected  by  the  parents,  has  had  little  or 
no  influence  on  the  differences  in  color  between  the  sexes.  It 
is  further  manifest,  in  all  the  cases  in  which  the  males  take 
exclusive  charge  of  the  nests  and  young,  that  the  destruction 
of  the  buighter-colored  males  would  be  far  more  influential  on 
the  character  of  the  race,  than  the  destruction  of  the  brighter- 
colored  females;  for  the  death  of  the  male  during  the  period  of 
incubation  or  nursing  would  entail  the  death  of  the  young,  so 
that  they  could  not  inherit  his  peculiarities;  yet,  in  many  of 
these  very  cases  the  males  are  more  conspicuously  colored  than 
the  females. 

In  most  of  the  Lophobranchii  (Pipe-fish,  Hippocampi,  &c.)  the 
males  have  either  marsupial  packs  or  hemispherical  depressions 
on  the  abdomen,  in  which  the  ova  laid  by  the  female  are  hatched. 
The  males  also  show  great  attachment  to  their  young.^^  The 
sexes  do  not  commonly  differ  much  in  color;  but  Dr.  Giinther 
believes  that  the  male  Plippocampi  are  rather  brighter  than  the 
females.  The  genus  Solenostoma,  however,  offers  a  curious  ex- 
ceptional case,*°  for  the  female  is  much  more  vividly-colored  and 
spotted  than  the  male,  and  she  alone  has  a  marsupial  sack  and 
hatches  the  eggs;  so  that  the  female  of  Solenostoma  differs  from 
all  the  other  Lophobranchii  in  this  latter  respect,  and  from  almost 
all  other  fishes,  in  being  more  brightly-colored  than  the  male.  It 
is  improbable  that  this  remarkable  double  inversion  of  character  in 
the  female  should  be  an  accidental  coincidence.  As  the  males  of 
several  fishes,  which  take  exclusive  charge  of  the  eggs  and  young, 
are  more  brightly-colored  than  the  females,  a,nd  as  here  the  fe- 
male Solenostoma  takes  the  same  charge  and  is  brighter  than  the 
male,  it  might  be  argued  that  the  conspicuous  colors  of  that  sex 
which  is  the  more  important  of  the  two  for  the  welfare  of  the 
offspring,  must  be  in   some  manner  protective.     But  from  the 

39  Yarrell,  'Hist,  of  British  Fishes,'  vol.  ii.  1836,  pp.  329,  338. 

*o  Dr.  Gunther,  since  publishing-  an  account  of  this  species  in  'The 
Fishes  of  Zanzibar,'  by  Col.  Playfair,  1866,  p.  137,  has  re-examined  the 
specimens,  and  has  given  me  the  above  information. 


FISHES.  343 

large  number  of  fishes,  of  which  the  males  are  either  permanently 
or  periodically  brighter  than  the  females,  but  whose  life  is  not  at 
all  more  important  for  the  welfare  of  the  species  than  that  of 
the  female,  this  view  can  hardly  be  maintained.  When  we  treat 
of  birds  we  shall  meet  with  analogous  cases,  where  there  has  been 
a  complete  inversion  of  the  usual  attributes  of  the  two  sexes,  and 
we  shall  then  give  what  appears  to  be  the  probable  explanation, 
namely,  that  the  males  have  selected  the  more  attractive  females, 
instead  of  the  latter  having  selected,  in  accordance  with  the  usual 
rule  throughout  the  animal  kingdom,  the  more  attractive  males. 

On  the  whole  we  may  conclude,  that  with  most  fishes,  in  which 
the  sexes  differ  in  color  or  in  other  ornamental  characters,  the 
males  originally  varied,  with  their  variations  transmitted  to  the 
same  sex,  and  accumulated  through  sexual  selection  by  attracting 
or  exciting  the  females.  In  many  cases,  however,  such  characters 
have  been  transferred,  either  partially  or  completely,  to  the  fe- 
males. In  other  cases,  again,  both  sexes  have  been  colored  alike 
for  the  sake  of  protection;  but  in  no  instance  does  it  appear  that 
the  female  alone  has  had  her  colors  or  other  characters  specially 
modified  for  this  latter  purpose. 

The  last  point  which  need  be  noticed  is  that  fishes  are  known 
to  make  various  noises,  some  of  which  are  described  as  being 
musical.  Dr.  Dufosse,  who  has  especially  attended  to  this  sub- 
ject, says  that  the  sounds  are  voluntarily  produced  in  several 
ways  by  different  fishes:  by  the  friction  of  the  pharyngeal  bones 
— by  the  vibration  of  certain  muscles  attacfied  to  the  swim-blad- 
der, which  serves  as  a  resounding-board — and  by  the  vibration  of 
the  intrinsic  muscles  of  the  swim-bladder.  By  this  latter  means 
the  Trigla  produces  pure  and  long-drawn  sounds  which  range 
over  nearly  an  octave.  But  the  most  interesting  case  for  us  is 
that  of  two  species  of  Ophidium,  in  which  the  males  alone  are 
provided  with  a  sound-producing  apparatus,  consisting  of  small 
movable  bones,  with  proper  muscles,  in  connection  with  the  swim- 
bladder.^^  The  drumming  of  the  Umbrinas  in  the  European  seas 
is  said  to  be  audible  from  a  depth  of  twenty  fathoms;  and  the 
fishermen  of  Rochelle  assert  "that  the  males  alone  make  the 
"noise  during  the  spawning-time;  and  that  it  is  possible  by  imi- 
"tating  it,  to  take  them  without  bait."*^  From  this  statement, 
and  more  especially  from  the  case  of  Ophidium,  it  is  almost  cer- 
tain that  in  this,  the  lowest  class  of  the  Vertebrata,  as  with  so 


41  'Comptes  Rendus.'  Tom.  xlvi,  1858,  p.  353.  Tom.  xlvii.  1858,  p.  916. 
Tom.  liv.  1862,  p.  393.  The  noise  made  by  the  Umbrinas  (Sciaena  aquila), 
is  said  by  some  authors  to  be  more  like  that  of  a  flute  or  org-an,  than 
drumming:  Br.  Zouteveen,  in  the  Dutch  translation  of  this  work  (vol. 
li.  p.  36),  gives  some  further  particulars  on  the  sounds  made  by  fishes. 

*2  The  Rev.  C.  Kingsley,  in  'Nature,'  May,  1S70,  p.  40. 


344  THE  DESCENT  OF  MAN. 

many  insects  and  spiders,  sound-producing  instruments  have,  at 
least  in  some  cases,  been  developed  through  sexual  selection,  as  a 
means  for  bringing  the  sexes  together. 

AMPHIBIANS. 

Urodela.  — I  will  begin  with  the  tailed  amphibians.  The  sexes 
of  salamanders  or  newts  often  differ  much  both  in  color  and 
structure.  In  some  species  prehensile  claws  are  developed  on 
the  fore-legs  of  the  males  during  the  breeding-season:  and  at 
this  season  in  the  male  Triton  palmipes  the  hind-feet  are  pro- 
vided with  a  swimming-web,  which  is  almost  completely  absorbed 
during  the  winter;    so  that  their  feet  then  resemble  those  of  the 


Fig.  32.  Triton  cristatus  (half  natural  size,  from  Bell's  'British  Rep- 
tiles'). Upper  figure,  male  during  tlie  breeding-season;  lower  fig- 
ure, female. 

female.*^  This  structure  no  doubt  aids  the  male  in  his  eager 
search  and  pursuit  of  the  female.  Whilst  courting  her  he  rapidly 
vibrates  the  end  of  his  tail.  With  our  common  newts  (Triton 
punctatus  and  cristatus)  a  deep,  much  Indented  crest  is  developed 
along  the  back  and  tail  of  the  male  during  the  breeding-season, 
which  disappears  during  the  winter.  Mr.  St.  George  Mivart  in- 
forms me  that  it  is  not  furnished  with  muscles,  and  thererore  can- 
not be  used  for  locomotion.  As  during  the  season  of  courtship  it 
becomes  edged  with  bright  colors,  there  can  hardly  be  a  doubt 
that  it  is  a  masculine  ornament.  In  many  species  the  body  presents 
strongly  contrasted,  though  lurid  tints,  and  these  become  more 
vivid  during  the  breeding-season.    The  male,  for  instance,  of  our 

*3Bell,   'History  of  British  Reptiles,'  2nd  edit.  1849,  pp.  156-159. 


AMPHIBIANS.  345 

common  little  newt  (Triton  punctatus)  is  "brownish-gray  above 
"passing  into  yellow  beneath,  which  in  the  spring  becomes  a  rich 
"bright  orange,  marked  everywhere  with  round  dark  spots."  The 
edge  of  the  crest  also  is  then  tipped  with  bright  red  or  violet. 
The  female  is  usually  of  a  yellowish-brown  color  with  scattered 
brown  dots,  and  the  lower  surface  is  often  quite  plain.**  The 
young  are  obscurely  tinted.  The  ova  are  fertilized  during  the  act 
of  deposition,  and  are  not  subsequently  tended  by  either  parent. 
We  may  therefore  conclude  that  the  males  have  acquired  their 
strongly-marked  colors  and  ornamental  appendages  through  sex- 
ual selection;  these  being  transmitted  either  to  the  male  offspring 
alone,  or  to  both  sexes. 

Anura  or  Batrachia. — With  many  frogs  and  toads  the  colors  evi- 
dently serve  as  a  protection,  such  as  the  bright  green  tints  of  tree- 
frogs  and  the  obscure  mottled  shades  of  many  terrestrial  species. 
The  most-conspicuously  colored  toad  which  I  ever  saw,  the  Phry- 
niscus  nigricans,*^  had  the  whole  upper  surface  of  the  body  as 
black  as  ink,  with  the  soles  of  the  feet  and  parts  of  the  abdomen 
spotted  with  the  brightest  vermilion.  It  crawled  about  the  bare, 
sandy  or  open  grassy  plains  of  La  Plata  under  a  scorching  sun, 
and  could  not  fail  to  catch  the  eye  of  every  passing  creature. 
These  colors  are  probably  beneficial  by  making  this  animal  known 
to  all  birds  of  prey  as  a  nauseous  mouthful. 

In  Nicaragua  there  is  a  little  frog  "dressed  in  a  bright  livery  of 
"red  and  blue"  which  does  not  conceal  itself  like  most  other 
species,  but  hops  about  during  the  daytime,  and  Mr.  Belt  says**' 
that  as  soon  as  he  saw  its  happy  sense  of  security,  he  felt  sure 
that  it  was  uneatable.  After  several  trials  he  succeeded  in  tempt- 
ing a  young  duck  to  snatch  up  a  young  one,  but  it  was  instantly 
rejected;  and  the  duck  "went  about  jerking  its  head,  as  if  trying 
"to  throw  off  some  unpleasant  taste." 

With  respect  to  sexual  differences  of  color.  Dr.  Giinther  does 
not  know  of  any  striking  instance  either  with  frogs  or  toads;  yet 
he  can  often  distinguish  the  male  from  the  female,  by  the  tints  of 
the  former  being  a  little  more  intense.  Nor  does  he  know  of  any 
striking  difference  in  external  structure  between  the  sexes,  ex- 
cepting the  prominences  which  become  developed  during  the 
breeding-season  on  the  front-legs  of  the  male,  by  which  he  is  en- 
abled to  hold  the  female.*^    It  is  surprising  that  these  animals 

**  Bell,  'History  of  British  Reptiles,'  2nd  edit.  1849,  pp.  146,  151. 

*5  'Zoology  of  the  Voyag-e  of  the  "Beagle,"  '  1843.    Bell,  ibid.  p.  49, 

*8  'The  Naturalist  in  Nicaragua,*  1874,  p.  321. 

*'7  The  male  alone  of  the  Bufo  sikimmensis  (Dr.  Anderson,  'Proc. 
Zoolog.  Soc'  1871,  p.  204)  has  two  plate-like  callosities  on  the  thorax 
and  certain  rugosities  on  the  fingers,  which  perhaps  subserve  the  same 
end  as  the  above-mention^  prominences. 


346  THE  DESCENT  OF  MAN. 

have  not  acquired  more  strongly-marked  sexual  characters;  for 
though  cold-blooded  their  passions  are  strong.  Dr.  Giinther  in- 
forms me  that  he  has  several  times  found  an  unfortunate  female 
toad  dead  and  smothered  from  having  been  so  closely  embraced 
by  three  or  four  males.  Progs  have  been  observed  by  Professor 
Hoffman  in  Giessen  fighting  all  day  long  during  the  breeding- 
season,  and  with  so  much  violence,  that  one  had  its  body  ripped 
open. 

Frogs  and  toads  offer  one  interesting  sexual  difference,  namely, 
in  the  musical  powers  possessed  by  the  males;  but  to  speak  of 
music,  when  applied  to  the  discordant  and  overwhelming  sounds 
emitted  by  male  bull-frogs  and  some  other  species,  seems,  ac- 
cording to  our  taste,  a  singularly  inappropriate  expression. 
Nevertheless,  certain  frogs  sing  in  a  decidedly  pleasing  manner. 
Near  Rio  Janeiro  I  used  often  to  sit  in  the  evening  to  listen  to  a 
number  of  little  Hylse,  perched  on  blades  of  grass  close  to  the  wa- 
ter, which  sent  forth  sweet  chirping  notes  in  harmony.  The  va- 
rious sounds  are  emitted  chiefly  by  the  males  during  the  breeding- 
season,  as  in  the  case  of  the  croaking  of  our  common  frog.*^  In 
accordance  with  this  fact  the  vocal  organs  of  the  males  are  more 
highly-developed  than  those  of  the  females.  In  some  genera  the 
males  alone  are  provided  with  sacs  which  open  into  the  larnyx.*^ 
For  instance,  in  the  edible  frog  (Rana  esculenta)  "the  sacs  are 
"peculiar  to  the  males,  and  become,  when  filled  with  air  in 
"the  act  of  croaking,  large  globular  bladders,  standing  out  one 
"on  each  side  of  the  head,  near  the  corners  of  the  mouth." 
The  croak  of  the  male  is  thus  rendered  exceedingly  powerful; 
whilst  that  of  the  female  is  only  a  slight  groaning  noise.^"  In  the 
several  genera  of  the  family  the  vocal  organs  differ  considerably 
in  structure,  and  their  development  in  all  cases  may  be  attributed 
to  sexual  selection. 

REPTILES. 

Chelonia. — Tortoises  and  turtles  do  not  offer  well-marKed  sexual 
differences.  In  some  species,  the  tail  of  the  male  is  longer  than 
that  of  the  female.  In  some,  the  plastron  or  lower  surface  of  the 
shell  of  the  male  is  slightly  concave  in  relation  to  the  back  of  the 
female.  The  male  of  the  mud-turtle  of  the  United  States  (Chrys- 
emys  picta)  has  claws  on  its  front-feet  twice  as  long  as  those  of 
the  female;  and  these  are  used  when  the  sexes  unite.^^  With  the 
huge  tortoise  of  the  Galapagos  Islands  (Testudo  nigra)  the  males 
are  said  to  grow  to  a  larger  size  than  the  females:  during  the 
pairing  season,  and  at  no  other  time,  the  male  utters  a  hoarse  bei- 

*8  Bell,  'History  of  British  Reptiles,'  1849,  p.  93. 

*»  J.  Bishop,  in  'Todd's  Cyclop,  of  Anat.  and  Phys.'  vol.  iv.  p.  1503. 

so  Bell,  ibid.  p.  112-114. 

51  'Mr,  C.  J.  Maynard,  'The  American  Naturalist,'  Dec.  1869,  p.  555. 


REPTILES.  347 

lowing  noise,  which  can  be  heard  at  the  distance  of  more  than  a 
hundred  yards;  the  female,  on  the  other  hand,  never  uses  her 
voice." 

With  the  Testudo  elegans  of  India,  it  is  said  "that  the  combats 
"of  the  males  may  be  heard  at  some  distance,  from  the  noise  they 
"produce  in  butting  against  each  other."^^ 

Croeodilia. — The  sexes  apparently  do  not  differ  in  color;  nor  do 
I  know  that  the  males  fight  together,  though  this  is  probable,  for 
some  kinds  make  a  prodigious  display  before  the  females.  Bar- 
tram-^*  describes  the  male  alligator  as  striving  to  win  the  female 
by  splashing  and  roaring  in  the  midst  of  a  lagoon,  "swollen  to  an 
"extent  ready  to  burst,  with  its  head  and  tail  lifted  up,  he  spins 
"or  twirls  round  on  the  surface  of  the  water,  like  an  Indian  chief 
"rehearsing  his  feats  of  war."  During  the  season  of  love,  a  musky 
odor  is  emitted  by  the  submaxillary  glands  of  the  crocodile,  and 
pervades  their  haunts.^^ 

Ophidia. — Dr.  Giinther  informs  me  that  the  males  are  always 
smaller  than  the  females,  and  generally  have  longer  and  slenderer 
tails;  but  he  knows  of  no  other  difference  in  external  structure. 
In  regard  to  color,  he  can  almost  always  distinguish  the  male  from 
the  female  by  his  more  strongly-pronounced  tints;  thus  the  black 
zigzag  band  on  the  back  of  the  male  English  viper  is  more  dis- 
tinctly defined  than  in  the  female.  The  difference  is  much  plainer 
in  the  rattle-snakes  of  N.  America,  the  male  of  which,  as  the 
keeper  in  the  Zoological  Gardens  showed  me,  can  at  once  be  dis- 
tinguished from  the  female  by  having  more  lurid  yellow  about  its 
whole  body.  In  S.  Africa  the  Bucephalus  capensis  presents  an 
analogous  difference,  for  the  female  "is  never  so  fully  variegated 
"with  yellow  on  the  sides  as  the  male."^®  The  male  of  the  Indian 
Dipsas  cynodon,  on  the  other  hand,  is  blackish-brown,  with  the 
belly  partly  black,  whilst  the  female  is  reddish  or  yellowish-olive, 
with  the  belly  either  uniform  yellowish  or  marbled  with  black. 
In  the  Tragops  dispar  of  the  same  country,  the  male  is  bright 
green,  and  the  female  bronze-colored."  No  doubt  the  colors  of 
some  snakes  are  protective,  as  shown  by  the  green  tints  of  tree- 
snakes,  and  the  various  mottled  shades  of  the  species  which  live  in 
sandy  places;  but  it  is  doubtful  whether  the  colors  of  many  kinds, 
for  instance  of  the  common  English  snake  and  viper,  serve  to 

52  See  my  'Journal  of  Researches  During-  the  Voyage  of  the  "Bea- 
"gle,"  '  1845,  p.  384. 

53  Dr.  Gunther,  'Reptiles  of  British  India,'  1864,  p.  7. 
^  'Travels  through  Carolina,'  &c.,  1791,  p.  128. 

=^  Owen,  'Anatomy  of  Vertebrates,'  vol.  i.  1866,  p=  615. 

58  Sir  Andrew  Smith,  'Zoolog.  of  S.  Africa:    Reptilia,'  1849,  pi.  s. 

"  Dr.  A.  Gunther,  'Reptiles  of  British  India,'  Ray  Soc.  1864,  pp.  304,  SOG. 


348  THE  DESCENT  OF  MAN. 

conceal  them;  and  this  is  still  more  doubtful  with  the  many  for- 
eign species  which  are  colored  with  extreme  elegance.  The  col- 
ors of  certain  species  are  very  different  in  the  adult  and  young 
snakes.*^^ 

During  the  breeding-season  the  anal  scent-glands  of  snakes  are 
in  active  function  ;^^  and  so  it  is  with  the  same  glands  in  lizards, 
and  as  we  have  seen  with  the  submaxillary  glands  of  crocodiles. 
As  the  males  of  most  animals  search  for  the  females,  these  odorif- 
erous glands  probably  serve  to  excite  or  charm  the  female,  rather 
than  to  guide  her  to  the  spot  where  the  male  may  be  found.  Male 
snakes,  though  appearing  so  sluggish,  are  amorous;  for  many 
have  been  observed  crowding  round  the  same  feihale,  and  even 
round  her  dead  body.  They  are  not  known  to  fight  together  from 
rivalry.  Their  intellectual  powers  are  higher  than  might  have 
been  anticipated.  In  the  Zoological  Gardens  they  soon  learn  not 
to  strike  at  the  iron  bar  with  which  their  cages  are  cleaned;  and 
Dr.  Keen  of  Philadelphia  informs  me  that  some  snakes  which  he 
kept,  learned  after  four  or  five  times  to  avoid  a  noose,  with  which 
they  were  at  first  easily  caught.  An  excellent  observer  in  Ceylon, 
Mr.  E.  Layard,  saw®°  a  cobra  thrust  its  head  through  a  narrow  hole 
and  swallow  a  toad.  "With  this  encumbrance  he  could  not  with- 
"draw  himself;  finding  this,  he  reluctantly  disgorged  the  precious 
"morsel,  which  began  to  move  off;  this  was  too  much  for  snake 
"philosophy  to  bear,  and  the  toad  was  again  seized,  and  again 
"was  the  snake,  after  violent  efforts  to  escape,  compelled  to  part 
"with  its  prey.  This  time,  however,  a  lesson  had  been  learnt, 
"and  the  toad  was  seized  by  one  leg,  withdrawn,  and  then  swal- 
"lowed  in  triurnph." 

The  keeper  in  the  Zoological  Gardens  is  positive  that  certain 
snakes,  for  instance  Crotalus  and  Python,  distinguish  him  from 
all  other  persons.  Cobras  kept  together  in  the  same  cage  appar- 
ently feel  some  attachment  towards  each  other.^^ 

It  does  not,  however,  follow  because  snakes  have  some  reason- 
ing power,  strong  passions  and  mutual  affection,  that  they  should 
likewise  be  endowed  with  sufficient  taste  to  admire  brilliant  colors 
in  their  partners,  so  as  to  lead  to  the  adornment  of  the  species 
through  sexual  selection.  Nevertheless,  it  is  difficult  to  account 
in  any  other  manner  for  the  extreme  beauty  of  certain  species; 
for  instance,  of  the  coral-snakes  of  S.  America,  which  are  of  a 
rich  red  with  black  and  yellow  transverse  bands.    I  well  remember 

58  Dr.  Stoliczka,  'Journal  of  Asiatic  Soc.  of  Bengral,'  vol.  xxxix.  1870, 
pp.  205,  211. 

B9  Owen,  'Anatomy  of  Vertebrates,'  vol.  i.  1866,  p.  615. 

«>  'Ilambles  in  Ceylon,'  in  'Annals  and  Mag.  of  Nat.  Hist.'  2nd  series. 
vol.  ix.  1852,  p.  333. 

«i  Dr.  Gunther,  'Reptiles  of  British  India,'  1864,  p.  340. 


REPTILES.  349 

how  much  surprise  I  felt  at  the  beauty  of  the  first  coral-snake 
which  I  saw  gliding  across  a  path  in  Brazil.  Snakes  colored  in 
this  peculiar  manner,  as  Mr.  Wallaee  states  on  the  authority  of 
Dr.  Giinther,^-  are  found  nowhere  else  in  the  world  except  in  S. 
America,  and  here  no  less  than  four  genera  occur.  One  of  these, 
Blaps,  is  venomous;  a  second  and  widely-distinct  genus  is  doubt- 
fully venomous,  and  the  two  others  are  quite  harmless.  The 
species  belonging  to  these  distinct  genera  inhabit  the  same  dis- 
tricts, and  are  so  like  each  other,  that  no  one  "but  a  naturalist 
"would  distinguish  the  harmless  from  the  poisonous  kinds." 
Hence,  as  Mr.  Wallace  believes,  the  innocuous  kinds  have  prob- 
ably acquired  their  colors  as  a  protection,  on  the  principle  of  imi- 
tation; for  they  would  naturally  be  thought  dangerous  by  their 
enemies.  The  cause,  however,  of  the  bright  colors  of  the  venom- 
ous Elaps  remains  to  be  explained,  and  this  may  perhaps  be  sexual 
selection. 

Snakes  produce  other  sounds  besides  hissing.  The  deadly 
Echis  carinata  has  on  its  sides  some  oblique  rows  of  scales  of  a  pe- 
culiar structure  with  serrated  edges;  and  when  this  snake  is  ex- 
cited, these  scales  are  rubbed  against  each  other,  which  produces 
"a  curious  prolonged,  almost  hissing  sound."^  With  respect  to 
the  rattling  of  the  rattle-snake,  we  have  at  last  some  definite  in- 
formation: for  Professor  Aughey  states,'''  that  on  two  occasions, 
being  himself  unseen,  he  watched  from  a  little  distance,  a  rattle- 
snake coiled  up  with  head  erect,  which  continued  to  rattle  at  short 
intervals  for  half  an  hour:  and  at  last  he  saw  another  snake 
approach,  and  when  they  met  they  paired.  Hence  he  is  satisfied 
that  one  of  the  uses  of  the  rattle  is  to  bring  the  sexes  together. 
Unfortunately  he  did  not  ascertain  whether  it  was  the  male  or 
the  female  which  remained  stationary  and  called  for  the  other. 
But  it  by  no  means  follows  from  the  above  fact  that  the  rattle  may 
not  be  of  use  to  these  snakes  in  other  ways,  as  a  warning  to  ani- 
mals which  would  otherwise  attack  them.  Nor  can  I  quite  dis- 
believe the  several  accounts  which  have  appeared  of  their  thus 
paralyzing  their  prey  with  fear.  Some  other  snakes  also  make  a 
distinct  noise  by  rapidly  vibrating  their  tails  against  the  surround- 
ing stalks  of  plants;  and  I  have  myself  heard  this  in  the  case  of  a 
Trigonocephalus  in  S.  America. 

Lacertilia. — The  males  of  some,  probably  of  many  kinds  of  liz- 
ards fight  together  from  rivalry.   Thus  the  arboreal  Anolis  crista- 
tellus  of  S.  America  is  extremely  pugnacious:  "During  the  spring 
"and  early  part  of  the  summer,  two  adult  males  rarely  meet  with- 
es -Westminster  Review,'  July  1st,  1867,  p.  32. 
«3  Dr.  Anderson,  'Proc.  Zoolog.  Soc'  1871,  p.  196. 
*  'The  American  Naturalist,'  1873,  p.  85. 


350  THE  DESCENT  OF  MAN. 

"out  a  contest.  On  first  seeing  one  another,  tliey  nod  their  heads 
"up  and  down  three  or  four  times,  and  at  the  same  time  expand- 
"ing  the  frill  or  pouch  beneath  the  throat;  their  eyes  glisten  with 
"rage,  and  after  waving  their  tails  from  side  to  side  for  a  few  sec- 
"onds,  as  if  to  gather  energy,  they  dart  at  each  other  furiously, 
"rolling  over  and  over,  and  holding  firmly  with  their  teeth.  The 
"conflict  generally  ends  in  one  of  the  combatants  losing  his  tail, 
"which  is  often  devoured  by  the  victor."  The  male  of  this  species 
is  considerably  larger  than  the  female;^'  and  this,  as  far  as  Dr. 
Gunther  has  been  able  to  ascertain,  is  the  general  rule  with  lizards 
of  all  kinds.  The  males  alone  of  the  Cyrtodactylus  rubidus  of  the 
Andaman  Islands  possesses  pre -anal  pores;  and  these  pores  judg- 
ing from  analogy  probably  serve  to  emit  an  odor.*^" 

The  sexes  often  differ  greatly  in  various  external  characters. 
The  male  of  the  above-mentioned  Anolis  is  furnished  with  a  crest 
which  runs  along  the  back  and  tail,  and  can  be  erected  at  pleas- 
ure; but  of  this  crest  the  female  does  not  exhibit  a  trace.  In  the 
Indian  Cophotis  ceylanica,  the  female  has  a  dorsal  crest,  though 
much  less  developed  than  in  the  male;  and  so  it  is,  as  Dr.  Gunther 
informs  me,  with  the  females  of  many  Iguanas,  Chameleons,  and 
other  lizards.  In  some  species,  however,  the  crest  is  equally 
developed  in  both  sexes,  as  in  the  Iguana  tuberculata.  In  the 
genus  Sitana,  the  males  alone  are  furnished  with  a  large  throat- 
pouch  (fig.  33),  which  can  be  folded  up  like  a  fan,  and  is  colored 
blue,  black,  and  red;  but  these  splendid  colors  are  exhibited  only 
during  the  pairing-season.  The  female  does  not  possess  even  a 
rudiment  of  this  appendage.  In  the  Anolis  cristatellus,  accord- 
ing to  Mr.  Austen,  the  throat  pouch,  which  is  bright  red  marbled 
with  yellow,  is  present  in  the  female,  though  in  a  rudimental  con- 
dition. Again,  in  certain  other  lizards,  both  sexes  are  equally 
well  provided  with  throat  pouches.  Here  we  see  with  species  be- 
longing to  the  same  group,  as  in  so  many  previous  cases,  the 
same  character  either  confined  to  the  males,  or  more  largely  de- 
veloped in  them  than  in  the  females,  or  again  equally  developed 
in  both  sexes.  The  little  lizards  of  the  genus  Draco,  which  glide 
through  the  air  on  their  rib-supported  parachutes,  and  which  in 
the  beauty  of  their  colors  baffle  description,  are  furnished  with 
skinny  appendages  to  the  throat  "like  the  wattles  of  gallinaceous 
birds."  These  become  erected  when  the  animal  is  excited.  They 
occur  in  both  sexes,  but  are  best  developed  when  the  male  arrives 
at  maturity,  at  which  age  the  middle  appendage  is  sometimes 
twice  as  long  as  the  head.    Most  of  the  species  likewise  have  a  low 

*»  Mr.  N.  Li.  Austen  kept  these  animals  alive  for  a  considerable  time; 
see  'Land  and  Water,'  July,  1S67,  p.  9. 
^^Stoliczka,  'Journal  of  Asiatic  Soc.  of  Bengal,'  vol.  xxxiv.  1870,  p.  166. 


REPTILES. 


3S1 


crest  running  along  the  neck;  and  this  is  much  more  developed  in 
the  full-grown  males,  than  in  the  females  or  young  males." 

A  Chinese  species  is  said  to  live  in  pairs  during  the  spring; 
"and  if  one  is  caught,  the  other  falls  from  the  tree  to  the  ground, 
"and  allows  itself  to  he  captured  with  impunity" — I  presume  from 
despair.''^ 

There  are  other  and  much  more  remarkable  differences  between 
the  sexes  of  certain  lizards.  The  male  of  Ceratophora  aspera 
bears  on  the  extremity  of  his  snout  an  appendage  half  as  long 


Fig.  33.  Sitana  minor.  Male  with 
the  g"ular  pouch  expanded 
(from  Gunther's  'Reptiles  of 
India'). 


Fig-.  34.  Ceratophora  Stod- 
dartii.  Upper  figure, 
male;  lower  figxire,  fe- 
male. 


as  the  head.  It  is  cylindrical,  covered  with  scales,  flexible,  and 
apparently  capable  of  erection:  in  the  female  it  is  quite  rudimen- 
tal.  In  a  second  species  of  the  same  genus  a  terminal  scale  forms 
a  minute  horn  on  the  summit  of  the  flexible  appendage-,  and  in  a 
third  species  (C.  Stoddartii,  fig.  34)  the  whole  appendage  is  con- 
verted into  a  horn,  which  is  usually  of  a  white  color,  but  assumes 
a  purplish  tint  when  the  animal  is  excited.  In  the  adult  male  of 
this  latter  species  the  horn  is  half  an  inch  in  length,  but  it  is  of 
quite  minute  size  in  the  female  and  in  the  young.  These  appen- 
dages, as  Dr.  Gtinther  has  remarked  to  me,  may  be  compared  with 
the  combs  of  gallinaceous  birds,  and  apparently  serve  as  orna- 
ments. 


«7  All  the  foregoing  statements  and  quotations,  in  regard  to  Cophotis, 
Sitana  and  Draco,  as  well  as  the  following  facts  in  regard  to  Cerato- 
phora and  Chamaeleon,  are  from  Dr.  Gunther  himself,  or  from  his 
magnificent  work  on  the  'Reptiles  of  British  India,'  Ray  See.  1864, 
pp.  122,  130,  135. 

«s  Mr.  Swinhoe,  'Proc.  Zoolog.  Soc'  1870,  p.  240. 


352 


THE  DESCENT  OF  MAN. 


In  the  genus  Chamgeleon  we  come  to  the  acme  of  difference  be- 
tween the  sexes.  The  upper  part  of  the  skull  of  the  male  C.  bifurcus 
(fig.  35),  an  inhabitant  of  Madagascar,  is  produced  into  two  great, 
solid,  bony  projections,  covered  with  scales  like  the  rest  of  the 
head;  and  of  this  wonderful  modification  of  structure  the  female 
exhibits  only  a  rudiment.  Again,  in  Chamseleon  Owenii  (fig.  36), 
from  the  West  Coast  of  Africa,  the  male  bears  on  his  snout  and 


Fig-.  35.   Chamaeleon  bifurcus.    Upper  figure,  male;  lower  figure,  female. 

forehead  three  curious  horns,  of  which  ihe  female  has  not  a  trace. 
These  horns  consist  of  an  excrescence  of  bone  covered  with  a 
smooth  sheath,  forming  part  of  the  general  integuments  of  the 
body,  so  that  they  are  identical  in  structure  with  those  of  a  bull, 
goat,  or  other  sheathhorned  ruminant.  Although  the  three  horns 
differ  so  much  in  appearance  from  the  two  great  prolongations  of 
the  skull  in  C.  bifurcus,  we  can  hardly  doubt  that  they  serve  the 
same  general  purpose  in  the  economy  of  these  two  animals.    The 


REPTILES. 


353 


first  conjecture,  which  will  occur  to  every  one,  is  that  they  are 
used  by  the  males  for  fighting  together;  and  as  these  animals  are 
very  quarrelsome,^  this  is  probably  a  correct  view.    Mr.  T.  W. 
Wood  also  informs  me  that  he  once  watched  two  individuals  of 
C.  pumilus,  fighting  violently  on  the  branch  of  a  tree;  they  flung 
their   heads   about   and 
tried  to  bite  each  other; 
they  then   rested   for   a 
time,     and     afterwards 
continued    their    battle. 
With     many     lizards, 
the  sexes  differ  slightly 
in  color,     the  tints  and 
stripes  of  the  males  be- 
ing   brighter   and    more 
distinctly    defined,    than 
in  the  females.  This,  for 
instance,     is     the    case 
with  the  above  Cophotis 
and   with   the  Acantho- 
dactylus   capensis   of   S. 
Africa.  In  a  Cordylus  of 
the    latter    country,   the 
male  is  either  much  red- 
der or  greener  than  the 
female.       In  the  Indian 
Calotes  nigrilabris  there  is  a  still  greater  difference;  the  lips  also 
of  the  male  are  black,  whilst  those  of  the  female  are  green.  In  our 
common  little  viviparous  lizard   (Zootoca  vivipara)   "the  under 
"side  of  the  body  and  base  of  the  tail  in  the  male  are  bright 
'^orange,  spotted  with  black;  in  the  female  these  parts  are  pale- 
"grayish-green  without  spots."'^    We  have  seen  that  the  males 
alone  of  Sltana  possess  a  throat-pouch;   and  this  is  splendidly 
tinted  with  blue,  black,  and  red.    In  the  Proctotretus  tenuis  of 
Chile  the  male  alone  is  marked  with  spots  of  blue,  green,  and  cop- 
pery-red.'^   In   many   6ases   the    males   retain   the    same    colors 
throughout  the  year,  but  in  others  they  become  much  brighter  dur- 
ing the  breeding-season;  I  may  give  as  an  additional  instance  the 
Calotes  maria,  which  at  this  season  has  a  bright  red  head,  the  rest 
of  the  body  being  green.^=* 


Fig-  36.    Chamaeleon   Owenii.    Upper 
figure,  male;  lower  figure,  female. 


69  Dr.  Bucholz,  'Monatsbericht  K.  Pieuss.  Akad.'  Jan.  1874,  p.  78. 

'°  Bell,  'History  of  British  Reptiles,'  2nd  edit.  1849,  p.  40. 

1  For  Proctotretus,  see  'Zoology  of  the  Voyage  of  the  "Beagle:"  Rep- 
tiles,' by  Mr.  Bell,  p.  8.  For  the  Lizards  of  S.  Africa,  see  'Zoology  of 
S.  Africa:  Reptiles,'  by  Sir  Andrew  Smith,  pi.  25  and  39.  For  the  In- 
dian Calotes,  see  'Reptiles  of  British  India,*  by  Dr.  Gunther,  p.  143, 

'2  Gunther  in  'Proc.  Zoolog.  Soc'  1870,  p.  778,  with  a  colored  figure. 
U 


354  THE  DESCENT  OF  MAN. 

Both  sexes  of  many  species  are  beautifully  colored  exactly  alike; 
and  there  is  no  reason  to  suppose  that  such  colors  are  protective. 
No  doubt  with  the  bright  green  kinds  which  live  in  the  midst  of 
vegetation,  this  color  serves  to  conceal  them,  and  in  N.  Patagonia 
I  saw  a  lizard  (Proctotretus  multimaculatus)  which,  when  fright- 
ened, flattened  its  body,  closed  its  eyes,  and  then  from  its  mottled 
tints  was  hardly  distinguishable  from  the  surrounding  sand.  But 
the  bright  colors  with  which  so  many  lizards  are  ornamented,  as 
well  as  their  various  curious  appendages,  were  probably  acquired 
by  the  males  as  an  attraction,  and  then  transmitted  either  to  their 
male  offspring  alone,  or  to  both  sexes.  Sexual  selection,  indeed, 
seems  to  have  played  almost  as  important  a  part  with  rep- 
tiles as  with  birds;  and  the  less  conspicuous  colors  of  the  females 
in  comparison  with  the  males  cannot  be  accounted  for,  as  Mr. 
Wallace  believes  to  be  the  case  with  birds,  by  the  greater  exposure 
of  the  females  to  danger  during  incubation. 


BIRDS.  355 


CHAPTER    XIII. 
SECONDARY    SEXUAL    CflARACTERS    OF    BIRDS. 

Sexual  differences— Law  of  battle— Special  weapons— Vocal  org-ans- In- 
strumental music— Love-antics  and  dances — Decorations,  permanent 
and  seasonal — Double  and  sing-le  annual  moults — Display  of  orna- 
ments by  the  males. 

Secondary  sexual  characters  are  more  diversified  and  conspic- 
uous in  birds,  though  not  perhaps  entailing  more  important 
changes  of  structure,  than  in  any  other  class  of  animals.  I  sTiall, 
therefore,  treat  the  subject  at  considerable  length.  Male  birds 
sometimes,  though  rarely  possess  special  weapons  for  fighting 
with  each  other.  They  charm  the  female  by  vocal  or  instrumental 
music  of  the  most  varied  kinds.  They  are  ornamented  by  all 
sorts  of  combs,  wattles,  protuberances,  horns,  air-distended  sacks, 
top-knots,  naked  shafts,  plumes  and  lengthened  feathers  grace- 
fully springing  from  all  parts  of  the  body.  The  beak  and  naked 
skin  about  the  head,  and  the  feathers  are  often  gorgeously  colored. 
The  males  sometimes  pay  their  court  by  dancing,  or  by  fantastic 
antics  performed  either  on  the  ground  or  in  the  air.  In  one  in- 
stance, at  least,  the  male  emits  a  musky  odor,  which  we  may  sup- 
pose serves  to  charm  or  excite  the  female;  for  that  excellent  ob- 
server, Mr.  Ramsay,^  says  of  the  Australian  muskduck  (Biziura 
lobata)  that  "the  smell  which  the  male  emits  during  the  summer 
"months  is  confined  to  that  sex,  and  in  some  individuals  is  re- 
"tained  throughout  the  year;  I  have  never,  even  in  the  breeding- 
"season,  shot  a  female  which  had  any  smell  of  musk."  So  power- 
ful is  this  odor  during  the  pairing-season,  that  it  can  be  detected 
long  before  the  bird  can  be  seen.^  On  the  whole,  birds  appear  to 
be  the  most  aesthetic  of  all  animals,  excepting  of  course  man,  and 
they  have  nearly  the  same  taste  for  the  beautiful  as  we  have.  This 
is  shown  by  our  enjoyment  of  the  singing  of  birds,  and  by  our 
women,  both  civilized  and  savage,  decking  their  heads  with  bor- 
rowed plumes,  and  using  gems  which  are  hardly  more  brilliantly 

3  'Ibis,'  vol.  iii.  (new  series)  1867,  p.  414. 

2  Gould,  'Handbook  to  the  Birds  of  Australia,'  1865,  vol.  ii.  p.  383. 


356  THE   DESCENT  OP  MAN. 

colored  than  the  naked  skin  and  wattles  of  certain  birds.  In  man, 
however,  when  cultivated,  the  sense  of  beauty  is  manifestly  a  far 
more  complex  feeling,  and  is  associated  with  various  intellectual 
ideas. 

Before  treating  of  the  sexual  characters  with  which  we  are  here 
more  particularly  concerned,  I  may  just  allude  to  certain  differ- 
ences between  the  sexes  which  apparently  depend  on  differences  in 
their  habits  of  life;  for  such  cases,  though  common  in  the  lower, 
are  rare  in  the  higher  classes.  Two  humming-birds  belonging  to 
the  genus  Eustephanus,  which  inhabit  the  island  of  Juan  Fernan- 
dez, were  long  thought  to  be  specifically  distinct,  but  are  now- 
known,  as  Mr.  Gould  informs  me,  to  be  the  male  and  female  of  the 
same  species,  and  they  differ  slightly  in  the  form  of  the  beak.  In 
another  genus  of  humming-birds  (Grypus),  the  beak  of  the  male 
is  serrated  along  the  margin  and  hooked  at  the  extremity,  thus 
differing  much  from  that  of  the  female.  In  the  Neomorpha  of 
New  Zealand,  there  is,  as  we  have  seen,  a  still  wider  difference  in 
the  form  of  the  beak  in  relation  to  the  manner  of  feeding  of  the 
two  sexes.  Something  of  the  same  kind  has  been  observed  with 
the  gold-finch  (Carduelis  elegans),  for  I  am  assured  by  Mr.  J. 
Jenner  Weir  that  the  birdcatchers  can  distinguish  the  males  by 
their  slightly  longer  beaks.  The  flocks  of  males  are  often  found 
feeding  on  the  seeds  of  the  teazle  (Dipsacus),  which  they  can 
reach  with  their  elongated  beaks,  whilst  the  females  more  com- 
monly feed  on  the  seeds  of  the  betony  or  Scrophularia.  With  a 
slight  difference  of  this  kind  as  a  foundation,  we  can  see  how  the 
beaks  of  the  two  sexes  might  be  made  to  differ  greatly  through 
natural  selection.  In  some  of  the  above  cases,  however,  it  is  pos- 
sible that  the  beaks  of  the  males  may  have  been  first  modified  in 
relation  to  their  contests  with  other  males;  and  that  this  after- 
wards led  to  slightly  changed  habits  of  life. 

Law  of  Battle.  — Almost  all  male  birds  are  extremely  pugnacious, 
using  their  beaks,  wings,  and  legs  for  fighting  together.  We  see 
this  every  spring  with  our  robins  and  sparrows.  The  smallest  of 
all  birds,  namely  the  humming-bird,  is  one  of  the  most  quarrel- 
some. Mr.  Gosse^  describes  a  battle  in  which  a  pair  seized  hold 
of  each  other's  beaks,  and  whirled  round  and  round,  till  they  al- 
most fell  to  the  ground;  and  M.  Montes  de  Oca,  in  speaking  of 
another  genus  of  humming-bird,  says  that  two  males  rarely  meet 
without  a  fierce  aerial  encounter:  when  kept  in  cages  "their  fight- 
"ing  has  mostly  ended  in  the  splitting  of  the  tongue  of  one  of  the 
"two,  which  then  surely  dies  from  being  unable  to  feed."^  With 
Waders,  the  males  of  the  common  water-hen  (Gallinula  chloropus) 

3  Quoted  by  Mr.  Gould,  'Introduction  to  the  Trochilldae,'  1861,  p.  29. 
*  Gould,  ibid.  p.  52, 


LAW    OF    BATTLE. 


357 


"when  pairing,  fight  violently  for  the  females:  they  stand  nearly 
"upright  in  the  water  and  strike  with  their  feet."  Two  were  seen 
to  be  thus  engaged  for  half  an  hour,  until  one  got  hold  of  the  head 
of  the  other,  which  would  have  been  killed,  ha#.  not  the  observer 
interfered;  the  female  all  the  time  looking  on  as  a  quiet  spectator.^ 
Mr.  Blyth  informs  me  that  the  males  of  an  allied  bird  (Gallicrex 
cristatus)  are  a  third  larger  than  the  females,  and  are  so  pug- 


Fig.  37.    The  Ruff  or  Machetes  pugnax  (from  Brehm's  'Thierleben'). 

nacious  during  the  breeding-season,  that  they  are  kept  by  the 
natives  of  Eastern  Bengal  for  the  sake  of  fighting.  Various  other 
birds  are  kept  in  India  for  the  same  purpose,  for  instance,  the 
bulbuls  (Pycnonotus  hasmorrhous)  which  "fight  with  great 
"spirit."® 

The  polygamous  ruff  (Machetes  pugnax,  fig.  37)  is  notorious  for 
his  extreme  pugnacity;  and  in  the  spring,  the  males,  which  are 
considerably  larger  than  the  females,  congregate  day  after  day  at 
a  particular  spot,  where  the  females  propose  to  lay  their  eggs. 
The  fowlers  discover  these  spots  by  the  turf  being  trampled  some- 
what bare.  Here  they  fight  very  much  like  gamecocks,  seizing 
each  other  with  their  beaks  and  striking  with  their  wings.    The 


^  W.  Thompson,  'Nat.  Hist  of  Ireland:    Birds,'  vol.  ii.  1850,  p.  327. 
^Jerdon,  'Birds  of  India,'  1863,  vol.  ii.  p.  96. 


24 


358  THE  DESCENT  OF  MAN, 

great  ruff  of  feathers  round  the  neck  is  then  erected,  and  accord- 
ing to  Col.  Montagu  "sweeps  the  ground  as  a  shield  to  defend  the 
"more  tender  parts;"  and  this  is  the  only  instance  known  to  me  in 
the  case  of  birds,  of  any  structure  serving  as  a  shield.  The  ruff 
of  feathers,  however,  from  its  varied  and  rich  colors  probably 
serves  in  chief  part  as  an  ornament.  Like  most  pugnacious  birds, 
they  seem  always  ready  to  fight,  and  when  closely  confined  often 
kill  each  other;  but  Montagu  observed  that  their  pugnacity  be- 
comes greater  during  the  spring,  when  the  long  feathers  on  their 
necks  are  fully  developed;  and  at  this  period  the  least  movement 
by  any  one  bird  provokes  a  general  battle.^  Of  the  pugnacity  of 
web-footed  birds,  two  instances  will  suffice:  in  Guiana  "bloody 
"fights  occur  during  the  breeding-season  between  the  males  of  the 
"wild  musk-duck  (Cairina  moschata) ;  and  where  these  fights  have 
"occurred  the  river  is  covered  for  some  distance  with  feathers."^ 
Birds  which  seem  ill-adapted  for  fighting  engage  in  fierce  conflicts; 
thus  the  stronger  males  of  the  pelican  drive  away  the  weaker 
ones,  snapping  with  their  huge  beaks  and  giving  heavy  blows 
with  their  wings.  Male  snipe  fight  together,  "tugging  and  push- 
"ing  each  other  with  their  bills  in  the  most  curious  manner  im- 
"aginable."  Some  few  birds  are  believed  never  to  fight;  this  is  the 
case,  according  to  Audubon,  with  one  of  the  woodpeckers  of  the 
United  States  (Picus  auratus),  although  "the  hens  are  followed  by 
even  half  a  dozen  of  their  gay  suitors."^ 

The  males  of  many  birds  are  larger  than  the  females,  and  this 
no  doubt  is  the  result  of  the  advantage  gained  by  the  larger  and 
stronger  males  over  their  rivals  during  many  generations.  The 
difference  in  size  between  the  two  sexes  is  carried  to  an  extreme 
point  in  several  Australian  species;  thus  the  male  musk-duck  (Bi- 
ziura)  and  the  male  Cincloramphus  cruralis  (allied  to  our  pipits) 
are  by  measurement  actually  twice  as  large  as  their  respective  fe- 
males.^°  With  many  other  birds  the  females  are  larger  than  the 
males;  and  as  formerly  remarked,  the  explanation  often  given, 
namely,  that  the  females  have  most  of  the  work  in  feeding  their 
young,  will  not  suffice.  In  some  few  cases,  as  we  shall  hereafter 
see,  the  females  apparently  have  acquired  their  greater  size  and 
strength  for  the  sake  of  conquering  other  females  and  obtaining 
possession  of  the  males. 

The  males  of  many  gallinaceous  birds,  especially  of  the  polyg- 
amous kinds,   are   furnished  with    special  weapons   for  fighting 


■7  Macgillivray,  'Hist.  Brit.  Birds,'  vol.  iv.  1852,  pp.  177-181. 

8  Sir  R.  Schomburgk,  in  'Journal  of  R.  Geograph.  Soc'  vol.  xiii.  1843, 
p.  31. 

0  'Ornithological  Biography,'  vol.  i.  p.  191.  For  pelicans  and  snipes, 
see  vol.  iii.  pp.  138,  477. 

10  Gould,  'Handbook  of  Birds  of  Australia,'  vol.  i.  p.  395;    vol.  ii.  p.  383. 


LAW    OF    BATTLE.  359 

with  their  rivals,  namelj''  spurs,  which  can  be  used  with  fearful 
effect.  It  has  been  recorded  by  a  trustworthy  writer^^  that  in 
Derbyshire  a  kite  struck  at  a  game-hen  accompanied  by  her  chick- 
ens, when  the  cock  rushed  to  the  rescue,  and  drove  his  spur 
right  through  the  eye  and  skull  of  the  aggressor.  The  spur  was 
with  difiBculty  drawn  from  the  skull,  and  as  the  kite  though  dead 
retained  his  grasp,  the  two  birds  were  firmly  locked  together;  but 
the  cock  when  disentangled  was  very  little  injured.  The  invin- 
cible courage  of  the  game-cock  is  notorious:  a  gentleman  who 
long  ago  witnessed  the  brutal  scene,  told  me  that  a  bird  had  both 
its  legs  broken  by  some  accident  in  the  cockpit,  and  the  owner  laid 
a  wager  that  if  the  legs  could  be  spliced  so  that  the  bird  could 
stand  upright,  he  would  continue  fighting.  This  was  effected  on 
the  spot,  and  the  bird  fought  with  undaunted  courage  until  he  re- 
ceived his  death-stroke.  In  Ceylon  a  closely  allied,  wild  species, 
the  Gallus  Stanleyi,  is  known  to  fight  desperately  "in  defense  of 
"his  seraglio,"  so  that  one  of  the  combatants  is  frequently  found 
dead.^-  An  Indian  partridge  (Ortygornis  gularis,)  the  male  of 
which  is  furnished  with  strong  and  sharp  spurs,  is  so  quarrelsome, 
"that  the  scars  of  former  fights  disfigure  the  breast  of  almost 
"every  bird  you  kill."^* 

The  males  of  almost  all  gallinaceous  birds,  even  those  which  are 
not  furnished  with  spurs,  engage  during  the  breeding-season  in 
fierce  conflicts.  The  Capercailzie  and  Black-cock  (Tetrao  urogallus 
and  T.  tetrix),  which  are  both  polygamists,  have  regular  ap- 
pointed places,  where  during  many  weeks  they  congregate  In 
numbers  to  fight  together  and  to  display  their  charms  before  the 
females.  Dr.  Y/.  Kovalevsky  informs  me  that  in  Russia  he  has 
seen  the  snow  all  bloody  on  the  arenas  where  the  capercailzie 
have  fought;  and  the  black-cocks  "make  the  feathers  fiy  in  every 
"direction,"  when  several  "engage  in  a  battle  royal."  The  elder 
Brehm  gives  a  curious  account  of  the  Balz,  as  the  love-dances  and 
love-songs  of  the  Black-cock  are  called  in  Germany.  The  bird 
utters  almost  continuously  the  strangest  noises:  "he  holds  his 
"tail  up  and  spreads  it  out  like  a  fan,  he  lifts  up  his  head  and  neck 
"with  all  the  feathers  erect,  and  stretches  his  wings  from  the 
"body.  Then  he  takes  a  fev/  jumps  in  different  directions,  some-- 
"times  in  a  circle,  and  presses  the  under  part  of  his  beak  so  hard 
"against  the  ground  that  the  chin  feathers  are  rubbed  off.  Dur- 
"ing  these  movements  he  beats  his  wings  and  turns  round  and 
"round.  The  more  ardent  he  grows  the  more  lively  he  becomes, 
"until  at  last  the  bird  appears  like  a  frantic  creature."  At  such 
times  the  black-cocks  are  so  absorbed  that  they  become  almost 

"■  Mr.  Hewitt,  in  the  'Poultry  Book  by  Teg-etmeier,'  1866,  p.  137. 
i^Layard,  'Annals  and  Mag.  of  Nat,  I-Iist.'  vol.  xiv.  1854,  p.  63, 
^3  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  574. 


360  THE  DESCENT  OF  MAN. 

blind  and  deaf,  but  less  so  than  the  capercailzie:  hence  bird  after 
bird  may  be  shot  on  the  same  spot,  or  even  caught  by  the  hand. 
After  performing  these  antics  the  males  begin  to  fight:  and  the 
same  black-cock,  in  order  to  prove  his  strength  over  several  an- 
tagonists, will  visit  in  the  course  of  one  morning  several  Balz- 
places,  which  remain  the  same  during  successive  years." 

The  peacock  with  his  long  train  appears  more  like  a  dandy  than 
a  warrior,  but  he  sometimes  engages  in  fierce  contests:  the  Rev. 
W.  Darwin  Fox  informs  me  that  at  some  little  distance  from  Ches- 
ter two  peacocks  became  so  excited  whilst  fighting,  that  they  flew 
over  the  whole  city,  still  engaged,  until  they  alighted  on  the  top 
of  St.  John's  tower. 

The  spur,  in  those  gallinaceous  birds  which  are  thus  provided, 
is  generally  single;  but  Polyplectron  (see  fig.  51,  p.  391)  has  two 
or  more  on  each  leg;  and  one  of  the  Blood-pheasants  (Ithaginis 
cruentus)  has  been  seen  with  five  spurs.  The  spurs  are  generally 
confined  to  the  male,  being  represented  by  mere  knobs  or  rudi- 
ments in  the  female;  but  the  females  of  the  Java  peacock  (Pavo 
muticus)  and,  as  I  am  informed  by  Mr.  Blyth,  of  the  small  fire- 
backed  pheasant  (Euplocamus  erythropthalmus)  possess  spurs. 
In  Galloperdix  it  is  usual  for  the  males  to  have  two  spurs,  and  for 
the  females  to  have  only  one  on  each  leg.^^  Hence  spurs  may 
be  considered  as  a  masculine  structure,  which  has  been  occasion- 
ally more  or  less  transferred  to  the  females.  Like  most  other  sec- 
ondary sexual  characters,  the  spurs  are  highly  variable,  both  in 
number  and  development,  in  the  same  species. 

Various  birds  have  spurs  on  their  wings.  But  the  Egyptian 
goose  (Chenalopex  segyptiaeus)  has  only  "bare  obtuse  knobs," 
and  these  probably  show  us  the  first  steps  by  which  true  spurs 
have  been  developed  in  other  species.  In  the  spur-winged  goose, 
Plectropterus  gambensis,  the  males  have  much  larger  spurs  than 
the  females;  and  they  use  them,  as  I  am  informed  by  Mr.  Bartlett, 
in  fighting  together,  so  that,  in  this  case,  the  wing-spurs  serve 
as  sexual  weapons;  but  according  to  Livingstone,  they  are  chiefly 
used  in  the  defense  of  the  young.  The  Palamedea  (fig.  38)  is 
armed  with  a  pair  of  spurs  on  each  wing;  and  these  are  such  for- 
midable weapons,  that  a  single  blow  has  been  known  to  drive  a 
dog  howling  away.  But  it  does  not  appear  that  the  spurs  in  this 
case,  or  in  that  of  some  of  the  spur-winged  rails  are  larger 
in    the    male     than     in    the    female.^^      In     certain     plovers, 


14  Brehm,  'Illust.  Thierleben,'  1867,  B.  iv.  s.  351.  Some  of  the  foregoing 
statements  are  taken  from  L.  Lloyd,  'The  Game  Birds  of  Sweden,*  &c., 
1867,  p.  79. 

13  Jerdon,  'Birds  of  India:  on  Ithaginis,'  vol.  iii.  p.  523;  on  Gallo- 
perdix, p.  541. 

16  For  the  Egyptian  goose,  see  Macgillivray,  'British  Birds,'  vol.  iv. 
p.   639.    For  Plectropterus,    'Livingstone's   Travels,'   p.   254.    For  Pala- 


LAW    OF    BATTLE. 


561 


however,  tke  wing-spurs  must  be  considered  as  a 
sexual  character.  Thus  in  the  male  of  our  common 
peewit  (Vanellus  cristatus)  the  tubercle  on  the  shoulder 
of  the  wing  becomes  more  prominent  during  the  breeding-season, 
and  the  males  fight  together.  In  some  species  of  Lobivanellus 
a  similar  tubercle  becomes  developed  during  the  breeding-season 
"into  a  short  horny  spur."  In  the  Australian  L.  lobatus  both  sexes 


Fig-.  38.    Palamedea  cornuta  (from  Brehm),  showing-  the  double  wing- 
spurs,  and  the  filament  on  the  head. 

have  spurs,  but  these  are  much  larger  in  the  males  than  in  the 
females.  In  an  allied  bird,  the  Hoplopterus  armatus,  the  spurs 
do  not  increase  in  size  during  the  breeding-season;  but  these  birds 
have  been  seen  in  Egypt  to  fight  together,  in  the  same  manner 
as  our  peewits,  by  turning  suddenly  in  the  air  and  striking  side- 


medea,    Brehm's   'Thierleben,'   B.    iv.    s.   740.     See,    also,    en   this    bird 
Azara,  'Voyag-es  dans  FAmerique  merid.'  torn.  iv.  1809,  pp.  179,  253. 


862  THE   DESCENT  OF  MAN. 

ways  at  each  other,  sometimes  with  fatal  results.  Thus  also  they 
drive  away  other  enemies.^" 

The  season  of  love  is  that  of  hattle;  but  the  males  of  some 
birds,  as  of  the  game-fowl  and  ruff,  and  even  the  young  males  of 
the  v/ild  turkey  and  grouse,^^  are  ready  to  fight  whenever  they 
meet.  The  presence  of  the  female  is  the  teterrima  belli  causa. 
The  Bengali  baboos  make  the  pretty  little  males  of  the  amadavat 
(Estrelda  amandava)  fight  together  by  placing  three  small  cages 
in  a  row,  with  a  female  in  the  middle;  after  a  little  time  the  two 
males  are  turned  loose,  and  immediately  a  desperate  battle  en- 
sues.^" When  many  males  congregate  at  the  same  appointed  spot 
and  fight  together,  as  in  the  case  of  grouse  and  various  other  birds, 
they  are  generally  attended  by  the  females,-"  which  afterwards 
pair  with  the  victorious  combatants.  But  in  some  cases  the  pair- 
ing precedes  instead  of  succeeding  the  combat:  thus  according  to 
Audubon,'^  several  males  of  the  Virginian  goat-sucker  (Caprimul- 
gus  virginianus)  "court  in  a  highly  entertaining  manner  the  fe- 
"male  and  no  sooner  has  she  made  her  choice,  than  her  approved 
"gives  chase  to  all  intruders,  and  drives  them  beyond  his  domin- 
"ions."  Generally  the  males  try  to  drive  away  or  kill  their  rivals 
before  they  pair.  It  does  not,  however,  appear  that  the  females 
invariably  prefer  the  victorious  males.  I  have  indeed  been  as- 
sured by  Dr.  W.  Kovalevsky  that  the  female  capercailzie  some- 
times steals  away  with  a  young  male  who  has  not  dared  to  enter 
the  arena  with  the  older  cocks,  in  the  same  manner  as  occasionally 
happens  with  the  does  of  the  red-deer  in  Scotland.  When  two 
males  contend  in  presence  of  a  single  female,  the  victor,  no  doubt, 
commonly  gains  his  desire;  but  some  of  these  battles  are  caused  by 
wandering  males  trying  to  distract  the  peace  of  an  already  mated 
pair.^^ 

Even  with  the  most  pugnacious  species  it  is  probable  that  the 

"  See,  on  our  peewit,  Mr.  R.  Carr  in  'Land  and  Water,*  Aug.  8th, 
1868,  p.  46.  In  regard  to  Lobivanellus,  see  Jerdon's  'Birds  of  India,' 
vol.  iii.  p.  647,  and  Gould's  'Handbook  of  Birds  of  Australia,'  vol.  ii. 
p.  220.    For  the  Holopterus,  see  Mr.  Allen  in  the  'Ibis,'  vol.  v.  1863,  p.  156. 

^8  Audubon,  'Ornith.  Biography,'  vol.  ii.  p.  492;  vol.  i.  pp.  4-13. 

19  Mr.  Blyth,  'Land  and  Water,'  1867,  p.  212. 

20  Richardson  on  Tetrao  umbellus  'Fauna  Bor.  Amer. :  Birds,'  1831, 
p.  343.  L.  Lloyd,  'Game  Birds  of  Sweden,'  1867,  pp.  22,  79,  on  the  caper- 
cailzie and  black-cock.  Brehm,  however,  asserts  ('Thierleben,'  &c., 
B.  iv.  s.  352)  that  in  Germany  the  g-ray-hens  do  not  g'enerally  attend 
the  Balzen  of  the  black-cocks,  but  this  is  an  exception  to  the  common 
rule;  possibly  the  liens  may  lie  hidden  in  the  surrounding-  busht>s,  as 
is  known  to  be  the  case  with  the  gray-hens  in  Scandinavia,  and  with 
other  species  in  N.  America. 

21  'Onithological  E'ography,'  vol.  ii.  p.  275. 

22  Brehm,  'Thierleben,'  &c.,  B.  iv.  1867,  p.  990.  Audubon,  'Ornith,  Bi- 
ography,' vol.  ii.  p.  492. 


VOCAL    MUSIC.  363 

pairing  does  not  depend  exclusively  on  the  mere  strength  and 
courage  of  the  male;  for  such  males  are  generally  decorated  with 
various  ornaments,  which  often  become  more  brilliant  during  the 
breeding-season,  and  which  are  sedulously  displayed  before  the 
females.  The  males  also  endeavor  to  charm  or  excite  their  mates 
by  love-notes,  songs,  and  antics;  and  the  courtship  is,  in  many  in- 
stances, a  prolonged  affair.  Hence  it  is  not  probable  that  the  fe- 
males are  indifferent  to  the  charms  of  the  opposite  sex,  or  that 
they  are  invariably  compelled  to  yield  to  the  victorious  males. 
It  is  more  probable  that  the  females  are  excited,  either  before  or 
after  the  conflict,  by  certain  males,  and  thus  unconsciously  prefer 
them.  In  the  case  of  Tetrao  umbellus,  a  good  observer-^  goes  so 
far  as  to  believe  that  the  battles  of  the  males  "are  all  a  sham,  per- 
"formed  to  show  themselves  to  the  greatest  advantage  before  the 
"admiring  females  who  assemble  around;  for  I  have  never  been 
"able  to  find  a  maimed  hero,  and  seldom  more  than  a  broken 
"feather."  I  shall  have  to  recur  to  this  subject,  but  I  may  here 
add  that  with  the  Tetrao  cupido  of  the  United  States,  about  a 
score  of  males  assemble  at  a  particular  spot,  and  strutting  about, 
make  the  whole  air  resound  with  their  extraordinary  noises.  At 
the  first  answer  from  a  female  the  males  begin  to  fight  furiously, 
and  the  weaker  give  way;  but  then,  according  to  Audubon,  both 
the  victors  and  vanquished  search  for  the  female,  so  that  the 
females  must  either  then  exert  a  choice,  or  the  battle  must  be  re- 
newed. So,  again,  with  one  of  the  field-starlings  of  the  United 
States  (Sturnella  ludoviciana)  the  males  engage  in  fierce  conflicts, 
"but  at  the  sight  of  a  female  they  ail  fly  after  her,  as  if  mad."^* 

Vocal  and  InstruTRental  Music. — With  birds  the  voice  serves  to 
express  various  emotions,  such  as  distress,  fear,  anger,  triumph,  or 
mere  happiness.  It  is  apparently  sometimes  used  to  excite  terror, 
as  in  the  ease  of  the  hissing  noise  made  by  some  nestling-birds. 
Audubon-^  relates  that  a  night-heron  (Ardea  nycticorax,  Linn.) 
which  he  kept  tame,  used  to  hide  itself  when  a  cat  approached, 
and  then  "suddenly  start  up  uttering  one  of  the  most  frightful 
"cries,  apparentl3'-  enjoying  the  cat's  alarm  and  flight."  The 
common  domestic  cock  clucks  to  the  hen,  and  the  hen  to  her 
chickens,  when  a  dainty  morsel  is  found.  The  hen,  when  she 
has  laid  an  q^^,  "repeats  the  same  note  very  often,  and  concludes 
"with  the  sixth  above,  which  she  holds  for  a  longer  time;"^^  and 
thus  she  expresses  her  joy.     Some  social  birds  apparently  call  to 

23  'Land  and  Water,'  July  25th,  1868,  p.  14. 

2*  Audubon's  'Ornitholog.  Biography;'    on  Tetrao  cupido,  vol.  ii.  p.  492; 
on  the  Sturnus,  vol.  ii.  p.  219. 
-^  'Ornithological  Biograph.'  vol.  v.  p.  601. 
2«  The  Hon.  Daines  Barrington,  'Philosoph.  Transact.'  1773,  p.  252. 


364  THE  DESCENT  OP  MAN. 

each  other  for  aid;  and  as  they  flit  from  tree  to  tree,  the  flock  is 
kept  together  by  chirp  answering  chirp.  During  the  nocturnal 
migrations  of  geese  and  other  water-fowl,  sonorous  clangs  from 
the  van  may  be  heard  in  the  darkness  overhead,  answered  by 
clangs  in  the  rear.  Certain  cries  serve  as  danger  signals,  which, 
as  the  sportsman  knows  to  his  cost,  are  understood  by  the  same 
species  and  by  others.  The  domestic  cock  crows,  and  the  hum- 
ming-bird chirps,  in  triumph  over  a  defeated  rival.  The  true 
song,  however,  of  most  birds  and  various  strange  cries  are  chiefly 
uttered  during  the  breeding-season,  and  serve  as  a  charm,  or 
merely  as  a  call-note,  to  the  other  sex. 

Naturalists  are  much  divided  with  respect  to  the  object  of  the 
singing  of  birds.  Few  more  careful  observers  ever  lived  than 
Montagu,  and  he  maintained  that  the  "males  of  song-birds  and  of 
"many  others  do  not  in  general  search  for  the  female,  but,  on  the 
"contrary,  their  business  in  the  spring  is  to  perch  on  some  con- 
"spicuous  spot,  breathing  out  their  full  and  amorous  notes,  which, 
"by  instinct,  the  female  knows,  and  repairs  to  the  spot  to  choose 
"her  mate."-^  Mr.  Jenner  Weir  informs  me  that  this  is  certainly 
the  case  with  the  nightingale.  Bechstein,  who  kept  birds  during 
his  whole  life,  asserts,  "that  the  female  canary  always  chooses  the 
"best  singer,  and  that  in  a  state  of  nature  the  female  finch  selects 
"that  male  out  of  a  hundred  whose  notes  please  her  most."-" 
There  can  be  no  doubt  that  birds  closely  attend  to  each  other's 
song.  Mr.  Weir  has  told  me  of  the  case  of  a  bullfinch  which 
had  been  taught  to  pipe  a  German  waltz,  and  who  was  so  good 
a  performer  that  he  cost  ten  guineas;  when  this  bird  was  first 
introduced  into  a  room  where  other  birds  were  kept  and  he  began 
to  sing,  all  the  others,  consisting  of  about  twenty  linnets  and 
canaries,  ranged  themselves  on  the  nearest  side  of  their  cages, 
and  listened  with  the  greatest  interest  to  the  new  performer. 
Many  naturalists  believe  that  the  singing  of  birds  is  almost  exclu- 
sively "the  effect  of  rivalry  and  emulation,"  and  not  for  the  sake 
of  charming  their  mates.  This  was  the  opinion  of  Daines  Bar- 
rington  and  White  of  Selborne,  who  both  especially  attended  to 
this  subject.^®  Barrington,  however,  admits  that  "superiority  in 
"song  gives  to  birds  an  amazing  ascendancy  over  others,  as  is 
"well  known  to  bird-catchers." 

It  is  certain  that  there  is  an  intense  degree  of  rivalry  between 
the  males  in  their  singing.  Bird-fanciers  match  their  birds  to 
see  which  will  sing  longest;    and  I  was  told  by  Mr.  Yarrell  that 


27  'Onithological  Dictionary.'  1833,  p.  475. 

28  'Naturg-eschichte  der  Stubenvogel,'  1840,  s.  4.  Mr.  Harrison  Weir 
likewise  writes  to  me:— "I  am  informed  that  the  best  singing  males 
"generally  get  a  mate  first,  when  they  are  bred  in  the  same  room." 

2»  'Philosophical  Transactions,'  1773,  p.  263.  White's  'Natural  History 
of  Selborne,'  1825,  vol.  i.  p.  246. 


VOCAL    MUSIC.  365 

a  first-rate  bird  will  sometimes  sing  till  lie  drops  down  almost 
dead,  or  according  to  Beclistein,^°  quite  dead  from  rupturing  a 
vessel  in  the  lungs.  Whatever  the  cause  may  be,  male  birds,  as 
I  hear  from  Mr.  Weir,  often  die  suddenly  during  the  season  of 
song.  That  the  habit  of  singing  is  sometimes  quite  independent 
of  love  is  clear,  for  a  sterile,  hybrid  canary-bird  has  been  de- 
scribed^^ as  singing  whilst  viewing  itself  in  a  mirror,  and  then 
dashing  at  its  own  image;  it  likewise  attacked  with  fury  a  fe- 
male canary,  when  put  into  the  same  cage.  The  jealousy  excited 
by  the  act  of  singing  is  constantly  taken  advantage  of  by  bird- 
catchers;  a  male,  in  good  song,  is  hidden  and  protected,  whilst  a 
stuffed  bird,  surrounded  by  limed  twigs,  is  exposed  to  view.  In 
this  manner,  as  Mr.  Weir  informs  me,  a  man  has  in  the  course  of 
a  single  day  caught  fifty,  and  in  one  instance  seventy,  male  chaf- 
finches. The  power  and  inclination  to  sing  differ  so  greatly  with 
birds  that  although  the  price  of  an  ordinary  male  chaffinch  is 
only  sixpence,  Mr.  Weir  saw  one  bird  for  which  the  bird-catcher 
asked  three  pounds;  the  test  of  a  really  good  singer  being  that 
it  will  continue  to  sing  whilst  the  cage  is  swung  round  the  owner's 
head. 

That  male  birds  should  sing  from  emulation  as  well  as  for 
charming  the  female,  is  not  at  all  incompatible;  and  it  might 
have  been  expected  that  these  two  habits  would  have  concurred, 
like  those  of  display  and  pugnacity.  Some  authors,  however, 
argue  that  the  song  of  the  male  cannot  serve  to  charm  the  female, 
because  the  females  of  some  few  species,  such  as  of  the  canary, 
robin,  lark,  and  bullfinch,  especially  when  in  a  state  of  widowhood, 
as  Bechstein  remarks,  pour  forth  fairly  melodious  strains.  In 
some  of  these  cases  the  habit  of  singing  may  be  in  part  attributed 
to  the  females  having  been  highly  fed  and  confined,'^-  for  this 
disturbs  all  the  usual  functions  connected  with  the  reproduction 
of  the  species.  Many  instances  have  already  been  given  of  the 
partial  transference  of  secondary  masculine  characters  to  the  fe- 
male, so  that  it  is  not  at  all  surprising  that  the  females  of  some 
species  should  possess  the  power  of  song.  It  has  also  been  argued, 
that  the  song  of  the  male  cannot  serve  as  a  charm,  because  the 
males  of  certain  species,  for  instance  of  the  robin,  sing  during 
the  autumn.^^  But  nothing  is  more  common  than  for  animals  to 
take  pleasure  in  practicing  whatever  instinct  they  follow  at  other 
times  for  some  real  good.  How  often  do  we  see  birds  which  fly 
easily,  gliding  and  sailing  through  the  air  obviously  for  pleasure? 


30  'Naturg-esch.  der  Stubenvogel,'   1840,  s.  252. 

=51  Mr.  Bold,  'Zoologist,'  1843-44,  p.  659. 

^-  D.  Barring-ton,  'Phil.  Transact.'  1773,  p.  262.  Bechstein,  'Stuben- 
vog-el,'  1840,  s.  4. 

33  This  is  likewise  the  case  with  the  water-ouzel,  see  Mr.  Hepburn  in 
the  'Zoologist,'  1845-1848,  p.  1068. 


366  THE  DESCENT  OF  MAN. 

The  cat  plays  with  the  captured  mouse,  and  the  cormorant  with 
the  captured  fish.  The  weaver-bird  (Ploceus),  when  confined  in  a 
cage,  amuses  itself  by  neatly  weaving  blades  of  grass  between 
the  wires  of  its  cage.  Birds  which  habitually  fight  during  the 
breeding-season  are  generally  ready  to  fight  at  all  times;  and 
the  males  of  the  capercailzie  sometimes  hold  their  Balzen  or  leks 
at  the  usual  place  of  assemblage  during  the  autumn."-*  Hence 
it  is  not  at  all  surprising  that  male  birds  should  continue  singing 
for  their  ov/n  amusement  after  the  season  for  courtship  is  over. 

As  shown  in  a  previous  chapter,  singing  is  to  a  certain  extent 
an  art,  and  is  much  improved  by  practice.  Birds  can  be  taught 
various  tunes,  and  even  the  unmelodious  sparrow  has  learnt  to 
sing  like  a  linnet.  They  acquire  the  song  of  their  foster  parents,=^^^ 
and  sometimes  that  of  their  neighbors.'''  All  the  common  song- 
sters belong  to  the  Order  of  Insessores,  and  their  vocal  organs  are 
much  more  complex  than  those  of  most  other  birds;  yet  it  is  a 
singular  fact  that  some  of  the  Insessores,  such  as  ravens,  crows, 
and  magpies,  possess  the  proper  apparatus,"'  though  they  never 
sing,  and  do  not  naturally  modulate  their  voices  to  any  great  ex- 
tent. Hunter  asserts^^  that  with  the  true  songsters  the  muscles 
of  the  larynx  are  stronger  in  the  males  than  in  the  females;  but 
with  this  slight  exception  there  is  no  difference  in  the  vocal  or- 
gans of  the  two  sexes,  although  the  males  of  most  species  sing 
so  much  better  and  more  continuously  than  the  females. 

It  is  remarkable  that  only  small  birds  properly  sing.  The 
Australian  genus  Menura,  however,  must  be  excepted;  for  the 
Menura  Alberti,  which  is  about  the  size  of  a  half-grown  turkey, 
not  only  mocks  other  birds,  but  "its  own  whistle  is  exceedingly 
"beautiful  and  varied."  The  males  congregate  and  form  "cor- 
roborying  places,"  where  they  sing,  raising  and  spreading  their 
tails  like  peacocks,  and  drooping  their  wings."^  It  is  also  remark- 
able that  birds  which  sing  well  are  rarely  decorated  with  brilliant 
colors  or  other  ornaments.  Of  our  British  birds,  excepting  the 
bullfinch  and  goldfinch,  the  best  songsters  are  plain-colored.  The 
kingfisher,  bee-eater,  roller,  hoopoe,  woodpeckers,  «&e.,  utter  harsh 
cries;    and  the  brilliant  birds  of  the  tropics  are  hardly  ever  song- 


3*  L.  Lloyd,   'Game  Birds  of  Sweden,'  1867,  p.  25. 

33  Barrington,  ibid.    p.  264.    Bechstein,  ibid.  s.  5. 

36  Bureau  de  la  Malle  gives  a  curious  instance  ('Annales  des  Sc. 
Nat.'  3rd  series,  Zoolog.  torn.  x.  p.  118)  of  some  wild  blackbirds  in  his 
garden  in  Paris  which  naturally  learnt  a  republican  air  from  a  caged 
bird. 

S7  Bishop,  in  'Todd's  Cyclop,  of  Anat.  and  Phys.'    vol.  iv.  p.  1496. 

38  As  stated  by  Barrington  in  'Philosoph.  Transact.'  1773,  p.  262. 

3»  Gould,  'Handbook  to  the  Birds  of  Australia,'  vol.  i.  1865,  pp.  308-310. 
See  also  Mr.  T.  W.  Wood  in  the  'Student,'  April  1870,  p.  125. 


VOCAL    MUSIC. 


367 


sters.**  Hence  bright  colors  and  the  power  of  song  seem  to  re- 
place each  other.  We  can  perceive  that  if  the  plumage  did  not 
vary  in  brightness,  or  if  bright  colors  were  dangerous  to  the  spe- 
cies, other  means  would  be  employed  to  charm  the  females;  and 
melody  of  voice  offers  one  such  means. 


ikdiMil^^ 


In  some  birds  the  vocal  organs  differ  greatly  in  the  two  sexes. 
In  the  Tetrao  cupido  (fig.  39)  the  male  has  two  bare,  orange- 
colored  sacks,  one  on  each  side  of  the  neck;  and  these  are  largely 
inflated  when  the  male,  during  the  breeding-season,  makes  his 
curious   hollow    sound,   audible    at   a   great   distance.    Audubon 


*"  See  remarks  to  this  effect  in  Gould's  'Introduction  to  the  Trochi* 
lidae,'  ISGl,  p.  22,, 


363  THE  DESCENT  OF  MAN. 

proved  that  the  sound  was  intimately  connected  with  this  appa- 
ratus (which  reminds  us  of  the  air-sacl£s  on  each  side  of  the  mouth 
of  certain  male  frogs),  for  he  found  that  the  sound  was  much 
diminished  when  one  of  the  sacks  of  a  tame  bird  was  pricked, 
and  when  both  were  pricked  it  was  altogether  stopped.  The 
female  has  "a  somewhat  similar,  though  smaller  naked  space  of 
"skin  on  the  neck;  but  this  is  not  capable  of  inflation."*^  The 
male  of  another  kind  of  grouse  (Tetrao  urophasianus),  whilst 
courting  the  female,  has  his  "bare  yellow  oesophagus  inflated  to  a 
"prodigious  size,  fully  half  as  large  as  the  body;"  and  he  then 
utters  various  grating,  deep,  hollow  tones.  With  his  neck-feath- 
ers erect,  his  wings  lowered,  and  buzzing  on  the  ground,  and 
his  long  pointed  tail  spread  out  like  a  fan,  he  displays  a  variety 
of  grotesque  attitudes.  The  oesophagus  of  the  female  is  not  in 
any  way  remarkable.*^ 

It  seems  now  well  made  out  that  the  great  throat  pouch  of 
the  European  male  bustard  (Otis  tarda),  and  of  at  least  four 
other  species,  does  not,  as  was  formerly  supposed,  serve  to  hold 
water,  but  is  connected  with  the  utterance  during  the  breeding- 
season  of  a  peculiar  sound  resembling  "ock."*'  A  crow-like  bird 
inhabiting  South  America  (Cephalopterus  ornatus,  fig.  40)  is 
called  the  umbrella-bird,  from  its  immense  top-knot,  formed  oS 
bare  white  quills  surmounted  by  dark  blue  plumes,  which  it  can 
elevate  into  a  great  dome  no  less  than  five  inches  in  diameter, 
covering  the  whole  head.  This  bird  has  on  its  neck  a  long,  thin, 
cylindrical  fleshy  appendage,  which  is  thickly  clothed  with  scale- 
like blue  feathers.  It  probably  serves  in  part  as  an  ornament, 
but  likewise  as  a  resounding  apparatus;  for  Mr.  Bates  found 
that  it  is  connected  "with  an  unusual  development  of  the  trachea 
"and  vocal  organs."  It  is  dilated  when  the  bird  utters  its  singu- 
larly deep,  loud  and  long  sustained  fluty  note.  The  head-crest 
and  neck-appendage  are  rudimentary  in  the  female.** 


41  'The  Sportsman  and  Naturalist  in  Canada,'  by  Major  W.  Ross 
King,  1866,  pp.  144-146.  Mr.  T.  W.  Wood  gives  in  the  'Student'  (April, 
1870,  p.  116)  an  excellent  account  of  the  attitude  and  habits  of  this 
bird  during  its  courtship.  He  states  that  the  ear-tufts  or  neck-plumes 
are  erected,  so  that  they  meet  over  the  crown  of  the  head.  See  his 
drawing,  fig.  39. 

*2  Richardson,  'Fauna  Bor.  American:  Birds,'  1831,  p.  359.  Audubon 
ibid.  vol.  iv.  p.  507. 

43  The  following  papers  have  been  lately  written  on  this  subject: 
Prof.  A.  Newton,  in  the  'Ibis,'  1862,  p.  107;  Dr.  Cullen,  ibid.  1865,  p.  145; 
Mr.  Flower,  in  'Proc.  Zool.  Soc'  1865,  p.  747;  and  Dr.  Murie,  in  'Proc. 
Zool.  Soc'  1868,  p.  471.  In  this  latter  paper  an  excellent  figure  is  given 
of  the  male  Australian  Bustard  in  full  display  with  the  sack  distended. 
It  is  a  singular  fact  that  the  sack  is  not  developed  in  all  the  males  of 
the  same  species. 

44  Bates,  'The  Naturalist  on  the  Amazons,'  1863,  vol.  ii.  p.  284;  Wal- 


VOCAL  MUSIC. 


369 


The  vocal  organs  of  various  web-footed  and  wading  birds  are 
extraordinarily  complex,  and  differ  to  a  certain  extent  in  the  two 
sexes.  In  some  cases  the  trachea  is  convoluted,  like  a  French 
horn,  and  is  deeply  embedded  in  the  sternum.    In  the  wild  swan 


Fig-.    40.    The    Umbrella-bird   or   Cephalopterus    ornatus    (male,    from 

Brehm). 

(Cygnus  ferus)  it  is  more  deeply  embedded  in  the  adult  male, 
than  in  the  adult  female  or  young  male.  In  the  male  Merganser 
the  enlarged  portion  of  the  trachea  is  furnished  with  an  additional 
pair  of  muscles.^  In  one  of  the  ducks,  however,  namely  Anas 
punctata,  the  bony  enlargement  is  only  a  little  more  developed 
in  the  male  than  in  the  female.^^  But  the  meaning  of  these  dif- 
ferences in  the  trachea  of  the  two  sexes  of  the  Anatidae  is  not 


lace,  in  'Proc.  Zool.  Soc'  1850,  p.  206.  A  new  species,  with  a  still  larger 
neck-appendag-e  (C.  pendulig-er),  has  lately  been  discovered,  siee  'Ibis,' 
vol.  i.  p.  457. 

*-5  Bishop,  in  Todd's  Cyclop,  of  Anat.  and  Phys.'  vol.  iv.  p.  1499. 

*»  Prof.  Newton,  'Proc.  Zoolog.  Soc'  1871,  p.  651 
25 


370  THE  DESCENT  OF  MAN. 

understood;  for  the  male  is  not  always  the  more  vociferous: 
thus  with  the  common  duck,  the  male  hisses,  whilst  the  female 
utters  a  loud  quack/^  In  both  sexes  of  one  of  the  cranes  (Grus 
Virgo)  the  trachea  penetrates  the  sternum,  but  presents  "certain 
"sexual  modifications."  In  the  male  of  the  black  stork  there  is 
also  a  well-marked  sexual  difference  in  the  length  and  curvature 
of  the  bronchi.^  Highly  important  structures  have,  therefore,  in 
these  cases  been  modified  according  to  sex. 

It  is  often  difficult  to  conjecture  whether  the  many  strange 
cries  and  notes  uttered  by  male  birds  during  the  breeding-season, 
serve  as  a  charm  or  merely  as  a  call  to  the  female.  The  soft 
cooing  of  the  turtle-dove  and  of  many  pigeons,  it  may  be  pre- 
sumed, pleases  the  female.  When  the  female  of  the  wild  turkey 
utters  her  call  in  the  morning,  the  male  answers  by  a  note  which 
differs  from  the  gobbling  noise  made,  when  with  erected  feathers, 
rustling  wings  and  distended  wattles,  he  puffs  and  struts  before 
her.^^  The  spel  of  the  black-cock  certainly  serves  as  a  call  to  the 
female,  for  it  has  been  known  to  bring  four  or  five  females  from 
a  distance  to  a  male  under  confinement;  but  as  the  black-cock 
continues  his  spel  for  hours  during  successive  days,  and  in  the 
case  of  the  capercailzie  "with  an  agony  of  passion,"  we  are  led 
to  suppose  that  the  females  which  are  present  are  thus  charmed.'" 
The  voice  of  the  common  rook  is  known  to  alter  during  the  breed- 
ing-season, and  is  therefore  in  some  way  sexual.^^  But  what  shall 
we  say  about  the  harsh  screams  of,  for  instance,  some  kinds  of 
macaws;  have  these  birds  as  bad  taste  for  musical  sounds  as  they 
apparently  have  for  color,  judging  by  the  inharmonious  con- 
trast of  their  bright  yellow  and  blue  plumage?  It  is  indeed  pos- 
sible that  without  any  advantage  being  thus  gained,  the  loud 
voices  of  many  male  birds  may  be  the  result  of  the  inherited 
effects  of  the  continued  use  of  their  vocal  organs,  when  excited 
by  the  strong  passions  of  love,  jealousy  and  rage;  but  to  this 
point  we  shall  recur  when  we  treat  of  quadrupeds. 

We  have  as  yet  spoken  only  of  the  voice,  but  the  males  of  vari- 
ous birds  practice,  during  their  courtship,  what  may  be  called 

*7  The  spoonbill  (Platalae)  has  its  trachea  convoluted  into  a  fignre 
of  eight,  and  yet  this  bird  (Jerdon,  'Birds  of  India,'  vol.  iii.  p.  763)  is 
mute;  but  Mr.  Blyth  informs  me  that  the  convolutions  are  not  con- 
stantly present,  so  that  perhaps  they  are  now  tending  towards 
abortion. 

*8  'Elements  of  Comp.  Anat.'  by  R.  Wagner,  Eng.  translat.  1845,  p. 
111.  With  respect  to  the  swan,  as  given  above,  Yarrell's  'Hist,  of  Brit- 
ish Birds,'  2nd  edit.  1S45,  vol.  iii.  p.  193. 

*9  C.  L.  Bonaparte,  quoted  in  the  'Naturalist  Library:  Birds,'  vol. 
xiv.  p.  126. 

50  L.  Lloyd,  'The  Game  Birds  of  Sweden,'  «S:c.,  1887,  pp.  22,  81. 

f^i  Jenner,  'Philosoph.  Transactions,'  1824,  p.  20. 


INSTRUMENTAL     MUSIC.  371 

instrumental  music.  Peacocks  and  Birds  of  Paradise  rattle  their 
quills  together.  Turkey-cocks  scrape  their  v/ings  against  the 
ground,  and  some  kinds  of  grouse  thus  produce  a  buzzing  sound. 
Another  North  American  grouse,  the  Tetrao  umbellus,  when  with 
his  tail  erect,  his  ruffs  displayed,  "he  shows  off  his  finery  to  the 
"females,  who  lie  hid  in  the  neighborhood,"  drums  by  rapidly 
striking  his  wings  together  above  his  back,  according  to  Mr.  R. 
Raymond,  and  not,  as  Audubon  thought,  by  striking  them  against 
his  sides.  The  sound  thus  produced  is  compared  by  some  to 
distant  thunder,  and  by  others  to  the  quick  roll  of  a  drum.  The 
female  never  drums,  "but  flies  directly  to  the  place  where  the 
"male  is  thus  engaged."  The  male  of  the  Kalij-pheasant,  in  the 
Himalayas,  "often  makes  a  singular  drumming  noise  with  his 
"wings,  not  unlike  the  sound  produced  by  shaking  a  stiff  piece  of 
"cloth."  On  the  west  coast  of  Africa  the  little  black-weavers 
(Ploceus?)  congregate  in  a  small  party  on  the  bushes  round  a 
small  open  space,  and  sing  and  glide  through  the  air  with  quiv- 
ering wings,  "which  make  a  rapid  whirring  sound  like  a  child's 
"rattle."  One  bird  after  another  thus  performs  for  hours  together, 
but  only  during  the  courting-season.  At  this  season  and  at  no 
other  time,  the  males  of  certain  night-jars  (Caprimulgus)  make 
a  strange  booming  noise  with  their  wings.  The  various  species 
of  wood-peckerp  strike  a  sonorous  branch  with  their  beaks,  with 
so  rapid  a  vibratory  movement  that  "the  head  appears  to  be  in 
"two  places  at  once."  The  sound  thus  produced  is  audible  at  a 
considerable  distance,  but  cannot  be  described;  and  I  feel  sure 
that  its  source  would  never  be  conjectured  by  any  one  hearing  it 
for  the  first  time.  As  this  jarring  sound  is  made  chiefly  during 
the  breeding-season,  it  has  been  considered  as  a  love-song;  but  it 
is  perhaps  more  strictly  a  love-call.  The  female,  when  driven 
from  her  nest,  has  been  observed  thus  to  call  her  m.ate,  who 
answered  in  the  same  manner  and  soon  appeared.  Lastly,  the 
male  Hoopoe  (Upupa  epops)  combines  vocal  and  instrumental 
music;  for  during  the  breeding-season  this  bird,  as  Mr.  Swinhoe 
observed,  first  draws  in  air,  and  then  taps  the  end  of  its  beak 
perpendicularly  down  against  a  stone  or  the  trunk  of  a  tree,  "when 
"the  breath  being  forced  down  the  tubular  bill  produces  the  cor- 
rect sound."  If  the  beak  is  not  thus  struck  against  some  object, 
the  sound  is  quite  different.  Air  is  at  the  same  time  swallowed, 
and  the  oesophagus  thus  becomes  much  swollen;  and  this  probably 
acts  as  a  resonator,  not  only  with  the  hoopoe,  but  with  pigeons 
and  other  birds.^- 


^-  For  the  foreg-oing  facts  see,  on  Birds  of  Paradise,  Brehm,  'Thier- 
lebeii,'  Band  iii.  s.  325.  On  Grouse,  Richardson,  'Fauna  Bor.  Americ. 
Birds,'  pp.  343  and  359;  Major  W.  Ross  King,  'The  Sportsman  in 
Canada,'  1866,  p.  156;  Mr.  Haymond,  in*  Prof.  Cox's  'Geol.  Survey  of  In- 


372 


THE  DESCENT  OF  MAN. 


In  the  foregoing  cases  sounds  are  made  by  the  aid  of  structures 
already  present  and  otherwise  necessary;  but  in  the  following 
cases  certain  feathers  have  been  specially  modified  for  the  express 
purpose  of  producing  sounds.  The  drumming,  bleating,  neigh- 
ing, or  thundering  noise  (as  expressed  by  different  observers) 
made  by  the  common  snipe  (Scolopax  gallinago)  must  have  sur- 
prised every  one  who  has  ever  heard  it.  This  bird,  during  the 
pairing-season,  flies  to  "perhaps  a  thousand  feet  in  height,"  and 
after  zig-zagging  about  for  a  time  descends  to  the  earth  in  a 
curved  line,  with  outspread  tail  and  quivering  pinions,  and  sur- 


Fig.  41.    Outer  tail  feather  of  Scolopax  gallinag-o  (from  'Proc.  Zool. 

SoC   1858). 

prising  velocity.  The  sound  is  emitted  only  during  this  rapid 
descent.  No  one  was  able  to  explain  the  cause,  until  M.  Meves 
observed  that  on  each  side  of  the  tail  the  outer  feathers  are 
peculiarly  formed  (fig.  41),  having  a  stiff  sabre-shaped  shaft,  with 


Fig".  42.    Outer  tail-feather  of  Scolo- 
pax frenata. 


Fig.  43.    Outer  tail-feather  of  Scolo- 
pax javensis. 


Fig.  44.  Primary  wing- 
feather  of  a  humming-bird, 
the  Selasphorus  platycercus 
(from  a  sketch  by  Mr.  Sal- 
vin).  Upper  figure,  that  of 
male;  lower  figure,  corres- 
ponding feather  of  female. 


the  oblique  barbs  of  unusual  length,  the  outer  webs  being  strongly 
bound  together.  He  found  that  by  blowing  on  these  feathers, 
or  by  fastening  them  to  a  long  thin  stick  and  waving  them  rap- 
idly through  the  air,  he  could  reproduce  the  drumming  noise 


diana,'  p.  227;  Audubon,  'American  Ornitholog.  Biograph.'  vol.  i.  p.  216. 
On  the  Kalij-pheasant,  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  533.  On  the 
Weavers,  'Livingstone's  Expedition  to  the  Zambesi,'  1865,  p.  425.  On 
Woodpeckers,  Macgillivray,  'Hist,  of  British  Birds,'  vol.  iii.  1840,  pp. 
84,  88,  89,  and  95.  On  the  Hoopoe  Mr.  Swinhoe,  in  Proc.  Zoolog.  Soc' 
June  23,  1863  and  1871,  p.  348.  On  the  Night-jar,  Audubon,  ibid.  vol.  ii. 
p.  255,  and  'American  Naturalist,'  1873,  p.  672.  The  English  Night-jar 
likewise  makes  in  the  spring  a  curious  noise  during  its  rapid  flight. 


■V'OCAL  AND   INSTRUMENTAL  MUSIC.  373 

made  by  the  living  bird.  Both  sexes  are  furnished  with  these 
feathers,  but  they  are  generally  larger  in  the  male  than  in  the 
female,  and  emit  a  deeper  note.  In  some  species,  as  in  S.  frenata 
(fig.  42),  four  feathers,  and  in  S.  javensis  (fig.  43),  no  less  than 
eight  on  each  side  of  the  tail  are  greatly  modified.  Different 
tones  are  emitted  by  the  feathers  of  the  different  species  when 
waved  through  the  air;  and  the  Scolopax  Wilsonii  of  the  United 
States  makes  a  switching  noise  whilst  descending  rapidly  to  the 
earth.°^ 

In  the  male  of  the  Chamsepetes  unicolor  (a  large  gallinaceous 
bird  of  America)  the  first  primary  wing-feather  is  arched  towards 
the  tip  and  is  much  more  attenuated  than  in  the  female.  In  an 
allied  bird,  the  Penelope  nigra,  Mr.  Salvin  observed  a  male,  which, 
whilst  it  flew  downwards  "with  outstretched  wings,  gave  forth 
"a  kind  of  crashing  rushing  noise,"  like  the  falling  of  a  tree.^* 
The  male  alone  of  one  of  the  Indian  bustards  (Sypheotides  auri- 
tus)  has  its  primary  wing-feathers  greatly  acuminated;  and  the 
male  of  an  allied  species  is  known  to  make  a  humming  no'ise 
whilst  courting  the  female."^  In  a  widely  different  group  of  birds, 
namely  Humming-birds,  the  males  alone  of  certain  kinds  have 
either  the  shafts  of  their  primary  wing-feathers  broadly  dilated, 
or  the  webs  abruptly  excised  towards  the  extremity.  The  male, 
for  instance,  of  Selasphorus  platycercus,  when  adult,  has  the  first 
primary  wing-feather  (fig.  44),  thus  excised.  Whilst  flying  from 
flower  to  flower,  he  makes  "a  shrill,  almost  whistling  noise  ;"^" 
but  it  did  not  appear  to  Mr.  Salvin  that  the  noise  was  intentionally 
made. 

Lastly,  in  several  species  of  a  sub-genus  of  Pipra  or  Manakin, 
the  males,  as  described  by  Mr.  Sclater,  have  their  secondary  wing- 
feathers  modified  in  a  still  more  remarkable  manner.  In  the 
brilliantly-colored  P.  deliciosa  the  first  three  secondaries  are 
thick-stemmed  and  curved  towards  the  body;  in  the  fourth  and 
fifth  (fig.  45,  a)  the  change  is  greater;  and  in  the  sixth  and 
seventh  (b,  c)  the  shaft  "is  thickened  to  an  extraordinary  degree, 
"forming  a  solid  horny  lump."  The  barbs  also  are  greatly 
changed  in  shape,  in  comparison  with  the  corresponding  feathers 


53  See  M.  Meves'  interesting-  paper  in  'Proc.  Zool.  Soc'  1858,  p.  199. 
For  the  habits  of  the  snipe,  Macgillivray,  'Hist,  British  Birds,'  vol.  iv. 
p.  371.  For  the  American  snipe,  Capt.  Blakiston,  'Ibis,'  vol.  v.  1863,  p. 
131. 

^*  Mr.  Salvin,  in  'Proc.  Zool.  Soc'  1867,  p.  160.  I  am  much  indebted 
to  this  distinguished  ornithologist  for  sketches  of  the  feathers  of  th© 
Chamaepetes,  and  for  other  information. 

55  Jerdon,  'Birds  of  India,'  vol.  iii.  pp.  618,  621. 

^  Gould,   'Introduction  to  the  Trochilidae,'  1861,  p.  49.    Salvin,   'Proc. 
Zoolog.  Soc'  1867,  p.  160. 
25 


374 


THE   DESCENT  OF  MAN. 


(d,  e,  f)  in  the  female.  Even  the  bones  of  the  wing,  which  sup- 
port these  singular  feathers  in  the  male,  are  said  by  Mr.  Eraser 
to  be  much  thickened.  These  little  birds  make  an  extraordinary 
noise,  the  first  "sharp  note  being  not  unlike  the  crack  of  a  whip.""' 


Fig.  45.  Secondary  wing-feathers  of  Pipra  deliciosa  (from  Mr.  Sclater, 
in  'Proc.  Zool.  Soc'  1860).  The  three  upper  feathers,  a,  b,  c,  from 
the  male;  the  three  lower  corresponding  feathers,  d,  e,  f,  from  the 
female. 

a  and  d,  fifth  secondary  wing-feather  of  male  and  female,  upper 
surface. 

b  and  e,  sixth  secondary,  upper  surface,  c  and  f,  seventh  secondary, 
lower  surface. 


57  Sclater,  in  'Proc.  Zool.  Soc'  1860,  p.  90,  and  in  'Ibis,'  vol.  iv.  1862,  p. 
175.    Also  Salvin,  in  'Ibis,'  1860,  p.  37. 


LOVE   ANTICS   AND  DANCES.  375 

The  diversity  of  the  sounds,  both  vocal  and  instrumental,  made 
by  the  males  of  many  birds  during  the  breeding-season,  and  the 
diversity  of  the  means  for  producing  such  sounds,  are  highly  re- 
markable. We  thus  gain  a  high  idea  of  their  importance  for 
sexual  purposes,  and  are  reminded  of  the  conclusion  arrived  at 
as  to  insects.  It  is  not  difficult  to  imagine  the  steps  by  which 
the  notes  of  a  bird,  primarily  used  as  a  mere  call  or  for  some 
other  purpose,  might  have  been  improved  into  a  melodious  love 
song.  In  the  case  of  the  modified  feathers,  by  which  the  drum- 
ming, whistling  or  roaring  noises  are  produced,  we  know  that 
some  birds  during  their  courtship  flutter,  shake,  or  rattle  their 
unmodified  feathers  together;  and  if  the  females  were  led  to 
select  the  best  performers,  the  males  which  possessed  the 
strongest  or  thickest,  or  most  attenuated  feathers,  situated 
on  any  part  of  the  body,  would  be  the  most  successful;  and 
thus  by  slow  degrees  the  feathers  might  be  modified  to  al- 
most any  extent.  The  females,  of  course,  would  not  notice 
each  slight  successive  alteration  in  shape,  but  only  the  sounds 
thus  produced.  It  is  a  curious  fact  that  in  the  same  class  of  ani- 
mals, sounds  so  different  as  the  drumming  of  the  snipe's  tail, 
the  tapping  of  the  woodpecker's  beak,  the  harsh  trumpet-like  cry 
of  certain  water-fowl,  the  cooing  of  the  turtle-dove,  and  the  song 
of  the  nightingale,  should  all  be  pleasing  to  the  females  of  the 
several  species.  But  we  must  not  judge  of  the  tastes  of  distinct 
species  by  a  uniform  standard;  nor  must  we  judge  by  the  standard 
of  man  s  taste.  Even  with  man,  we  should  remember  what  dis- 
cordant noises,  the  beating  of  tom-toms  and  the  shrill  notes  of 
reeds,  please  the  ears  of  savages.  Sir  S.  Baker  remarks,^^  that 
"as  the  stomach  of  the  Arab  prefers  the  raw  meat  and  reeking 
"liver  taken  hot  from  the  animal,  so  does  his  ear  prefer  his 
"equally  coarse  and  discordant  music  to  all  other." 

Love-Antics  and  Dances. — The  curious  love  gestures  of  some 
birds  have  already  been  incidentally  noticed;  so  that  little  need 
here  be  added.  In  Northern  America,  large  numbers  of  a  grouse, 
the  Tetrao  phasianellus,  meet  every  morning  during  the  breeding- 
season  on  a  selected  level  spot,  and  here  they  run  round  and 
round  in  a  circle  of  about  fifteen  or  twenty  feet  in  diameter,  so 
that  the  ground  is  worn  quite  bare,  like  a  fairy-ring.  In  these 
Partridge-dances,  as  they  are  called  by  the  hunters,  the  birds 
assume  the  strangest  attitudes,  and  run  round,  some  to  the  left 
and  some  to  the  right.  Audubon  describes  the  males  of  a  heron 
(Ardea  herodias)  as  walking  about  on  their  long  legs  with  great 
dignity  before  the  females,  bidding  defiance  to  their  rivals.  With 
one  of  the  disgusting  carrion-vultures  (Cathartes  jota)  the  same 

58  The  Nile  Tributaries  of  Abyssinia,'   1867,  p.  203. 


376  THE  DESCENT  OF  MAN. 

naturalist  states  that  "the  gesticulations  and  parade  of  the  males 
"at  the  beginning  of  the  love-season  are  extremely  ludicrous." 
Certain  birds  perform  their  love  antics  on  the  wing,  as  we  have 
seen  with  the  black  African  weaver,  instead  of  on  the  ground. 
During  the  spring  our  little  white  throat  (Sylvia  cinerea)  often 
rises  a  few  feet  or  yards  in  the  air  above  some  bush,  and  "flut- 
"ters  with  a  fitful  and  fantastic  motion,  singing  all  the  while, 
"and  then  drops  to  its  perch."  The  great  English  bustard  throws 
himself  into  indescribably  odd  attitudes  whilst  courting  the  fe- 
male, as  has  been  figured  by  Wolf.  An  allied  Indian  bustard 
(Otis  bengalensis)  at  such  times  "rises  perpendicularly  into  the 
"air  with  a  hurried  flapping  of  his  wings,  raising  his  crest  and 
"puffing  out  the  feathers  of  his  neck  and  breast,  and  then  drops 
"to  the  ground;"  he  repeats  this  maneuver  several  times,  at  the 
same  time  humming  in  a  peculiar  tone.  Such  females  as  happen 
to  be  near  "obey  this  saltatory  summons,"  and  when  they  approach 
he  trails  his  wings  and  spreads  his  tail  like  a  turkey-cock.^^ 

But  the  most  curious  case  is  afforded  by  three  allied  genera  of 
Australian  birds,  the  famous  Bower-birds, — no  doubt  the  co- 
descendants  of  some  ancient  species  which  first  acquired  the 
strange  instinct  of  constructing  bowers  for  performing  their  love- 
antics.  The  bowers  (fig.  46),  which,  as  we  shall  hereafter  see,  are 
decorated  with  feathers,  shells,  bones,  and  leaves,  are  built  on 
the  ground  for  the  sole  purpose  of  courtship,  for  their  nests  are 
formed  in  trees.  Both  sexes  assist  in  the  erection  of  the  bowers, 
but  the  male  is  the  principal  workman.  So  strong  is  this  instinct 
that  it  is  practiced  under  confinement,  and  Mr.  Strange  has  de- 
scribed^" the  habits  of  some  Satin  Bower-birds  which  he  kept  in 
an  aviary  in  New  South  Wales.  "At  times  the  male  will  chase 
"the  female  all  over  the  aviary,  then  go  to  the  bower,  pick  up  a 
"gay  feather  or  a  large  leaf,  utter  a  curious  kind  of  a  note,  set  all 
"his  feathers  erect,  run  round  the  bower  and  become  so  excited 
"that  his  eyes  appear  ready  to  start  from  his  head;  he  continues 
"opening  first  one  wing  then  the  other,  uttering  a  low,  whistling 
"note,  and,  like  the  domestic  cock,  seems  to  be  picking  up  some- 
"thing  from  the  ground  until  at  last  the  female  goes  gently 
"towards   him."    Captain   Stokes  has   described  the  habits   and 


69  For  Tetrao  phasianellus,  see  Richardson,  'Fauna,  Bor.  America,' 
p.  361,  and  for  further  particulars  Capt.  Blakiston,  'Ibis,'  1863,  p.  125. 
For  the  Cathartes  and  Ardea,  Audubon,  'Ornith.  Biography,'  vol.  ii. 
p.  51,  and  vol.  iii.  p.  89,  On  the  White-throat,  Macgillivray,  'Hist.  Brit- 
ish Birds,'  vol.  ii.  p.  354.  On  the  Indian  Bustard,  Jerdon,  'Birds  of 
India,  vol.  ill.  p.  618. 

80  Gould,  'Handbook  to  the  Birds  of  Australia,'  vol.  i.  pp.  444,  449, 
455.  The  bower  of  the  Satin  Bower-bird  may  be  seen  in  the  Zoological 
Society's  Gardens,  Regent's  Park. 


DECORATION.  377 

"play-houses"  of  another  species,  the  Great  Bower-bird,  which 
was  seen  "amusing  itself  by  flying  backwards  and  forwards,  taking 
"a  shell  alternately  from  each  side,  and  carrying  it  through  the 
"archway  in  its  mouth."  These  curious  structures,  formed  solely 
as  halls  of  assemblage,  where  both  sexes  amuse  themselves  and 
pay  their  court,  must  cost  the  birds  m^uch  labor.  The  bower,  for 
instance,  of  the  Fawn-breasted  species,  is  nearly  four  feet  in 
length,  eighteen  inches  in  height,  and  is  raised  on  a  thick  platform 
of  sticks. 

Decoration.  — I  will  first  discuss  the  cases  in  which  the  males 
are  ornamented  either  exclusively  or  in  a  much  higher  degree 
than  the  females,  and  in  a  succeeding  chapter  those  in  which 
both  sexes  are  equally  ornamented,  and  finally  the  rare  cases  in 
which  the  female  is  somewhat  more  brightly-colored  than  the 
male.  As  with  the  artificial  ornaments  used  by  savage  and 
civilized  men,  so  with  the  natural  ornaments  of  birds,  the  head 
is  the  chief  seat  of  decoration."^  The  ornaments,  as  mentioned 
at  the  commencement  of  this  chapter,  are  wonderfully  diver- 
sified. The  plumes  on  the  front  or  back  of  the  head  consist  of 
variously-shaped  feathers,  sometimes  capable  of  erection  or  expan- 
sion, by  which  their  beautiful  colors  are  fully  displayed.  Ele- 
gant ear-tufts  (see  fig.  39,  ante)  are  occasionally  present.  The 
head  is  sometimes  covered  with  velvety  down,  as  with  the  pheas- 
ant; or  is  naked  and  vividly  colored.  The  throat,  also,  is  some- 
times ornamented  with  a  beard,  wattles,  or  caruncles.  Such  ap- 
pendages are  generally  brightly-colored,  and  no  doubt  serve  as 
ornaments,  though  not  always  ornamental  in  our  eyes;  for  whilst 
the  male  is  in  the  act  of  courting  the  female,  they  often  swell 
and  assume  vivid  tints,  as  in  the  male  turkey.  At  such  times 
the  fleshy  appendages  about  the  head  of  the  male  Tragopan  pheas- 
ant (Ceriornis  Temminckii)  swell  into  a  large  lappet  on  the 
throat  and  into  two  horns,  one  on  each  side  of  the  splendid  top- 
knot; and  these  are  then  colored  of  the  most  intense  blue  which 
I  have  ever  beheld. ^^  The  African  hornbill  (Bucorax  abyssinicus) 
inflates  the  scarlet  bladder-like  wattle  on  its  neck,  and  with  its 
wings  drooping  and  tail  expanded  "makes  quite  a  grand  appear- 
"ance."^  Even  the  iris  of  the  eye  is  sometimes  more  brightly- 
colored  in  the  male  than  in  the  female;  and  this  is  frequently  the 
case  with  the  beak,  for  instance,  in  our  common  blackbird.  In 
Buceros  corrugatus,  the  whole  beak  and  immense  casque  are  col- 

61  See  remarks  to  this  effect,  on  the  'Feeling  of  Beauty  among  Ani- 
mals,' by  Mr.  J.  Shaw,  in  the  'Athenaeum,'  Nov.  24th,  1866,  p.  681. 

«2  See  Dr.  Murie's  account  with  colored  figures  in  'Proc.  Zoolog.  Soc' 
1872,  p.  730. 

^3  Mr.  Monteiro,  'Ibis,'  vol.  iv.  1862,  p.  339. 


378 


THE  DESCENT  OF  MAN. 


ored  more  conspicuously  in  the  niale  than  in  the  female;  and  "the 
"oblique  grooves  upon  the  sides  of  the  lower  mandible  are  pe- 
"culiar  to  the  male  sex."" 


Fig.  46.  Bower-bird,  Chlamydera  maculata,  with  bower  (from  Brehm). 

i 
The  head,  again,  often  supports  fleshy  appendages,  filaments, 
and  solid  protuberances.  These,  if  not  common  to  both  sexes, 
are  always  confined  to  the  males.  The  solid  protuberances  have 
been  described  in  detail  by  Dr.  W.  Marshall,''^  who  shows  that 
they  are  formed  either  of  cancellated  bone  coated  with  skin,  or 
of  dermal  and  other  tissues.  With  mammals  true  horns  are 
always  supported  on  the  frontal  bones,  but  with  birds  various 
bones  have  been  modified  for  this  purpose;    and  in  species  of  the 


8*  'Land  and  Water,'  1868,  p.  217. 

«•''  Ueber  die  Schadelhocker,'  &c.,  'Niederlandischen  Archjv  fur  Zoo- 
logie,'  B.  I.  Heft,  2,  1872. 


DECORATION.  379 

same  group  the  protuberances  may  have  cores  of  bone,  or  be 
quite  destitute  of  them,  with  intermediate  gradations  connecting 
these  two  extremes.  Hence,  as  Dr.  Marshall  justly  remarks, 
variations  of  the  most  different  kinds  have  served  for  the  de- 
velopment through  sexual  selection  of  these  ornamental  appen- 
dages. Elongated  feathers  or  plumes  spring  from  almost  every 
part  of  the  body.  The  feathers  on  the  throat  and  breast  are 
sometimes  developed  into  beautiful  ruffs  and  collars.  The  tail- 
feathers  are  frequently  increased  in  length;  as  we  see  in  the 
tail-coverts  of  the  peacock,  and  in  the  tail  itself  of  the  Argus 
pheasant.  With  the  peacock  even  the  bones  of  the  tail  have 
been  modified  to  support  the  heavy  tail-coverts.'^^  The  body  of 
the  Argus  is  not  larger  than  that  of  a  fowl;  yet  the  length  from 
the  end  of  the  beak  to  the  extremity  of  the  tail  is  no  less  than 
five  feet  three  inches,"'  and  that  of  the  beautifully  ocellated  sec- 
ondary wing-feathers  nearly  three  feet.  In  a  small  African  night- 
jar (Cosmetornis  vexillarius)  one  of  the  primary  wing-feathers, 
during  the  breeding-season,  attains  a  length  of  twenty-six  inches, 
whilst  the  bird  itself  is  only  ten  inches  in  length.  In  another 
closely-allied  genus  of  night-jars,  the  shafts  of  the  elongated  wing- 
feathers  are  naked,  except  at  the  extremity,  where  there  is  a 
disc.*'*  Again,  in  another  genus  of  night-jars,  the  tail-feathers 
are  even  still  more  prodigiously  developed.  In  general  the  feath- 
ers of  the  tail  are  more  often  elongated  than  those  of  the  wings, 
as  any  great  elongation  of  the  latter  impedes  flight.  We  thus  see 
that  in  closely-allied  birds  ornaments  of  the  same  kind  have  been 
gained  by  the  males  through  the  development  of  widely  different 
feathers. 

It  is  a  curious  fact  that  the  feathers  of  species  belonging  to 
very  distinct  groups  have  been  modified  in  almost  exactly  the 
same  peculiar  manner.  Thus  the  wing-feathers  in  one  of  the 
above-mentioned  night-jars  are  bare  along  the  shaft,  and  ter- 
minate in  a  disc;  or  are,  as  they  are  sometimes  called,  spoon  or 
racket-shaped.  Feathers  of  this  kind  occur  in  the  tail  of  a 
motmot  (Eumomota  superciliaris),  of  a  king-fisher,  finch,  hum- 
ming-bird, parrot,  several  Indian  drongos  (Dicrurus  and  Edolius, 
in  one  of  which  the  disc  stands  vertically),  and  in  the  tail  of 
certain  birds  of  paradise.  In  these  latter  birds,  similar  feathers, 
beautifully  ocellated,  ornament  the  head,  as  is  likewise  the  case 
with  some  gallinaceous  birds.  In  an  Indian  bustard  (Sypheotides 
auritus)  the  feathers  forming  the  ear-tufts,  which  are  about  four 
Inches  in  length,  also  terminate  in  discs.^^    It  is  a  most  singular 


««  Dr.  W.  Marshall,  'Tiber  den  Vogelschwanz,'  ibid.    B.  I.  Heft  2,  1872, 
«'^  Jardine's  'Naturalist  Library:  Birds,'  vol.  xiv.  p.  166. 
^  Sclater,  in  the  'Ibis,'  vol.  vi.  1864,  p.  114.    Livingstone,  'Expedition 
to  the  Zambesi,'  1865,  p.  66. 
«8  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  620. 


380  THE  DESCENT  OF  MAN. 

fact  that  the  motmots,  as  Mr.  Saivin  has  clearly  shown,™  give  to 
their  tail-feathers  the  racket-shape  by  biting  ofE  the  barbs,  and, 
further,  that  this  continued  mutilation  has  produced  a  certain 
amount  of  inherited  effect. 

Again  the  barbs  of  the  feathers  in  various  widely-distinct 
birds  are  filamentous  or  plumose,  as  with  some  herons,  ibises, 
birds  of  paradise,  and  Gallinacese.  In  other  cases  the  barbs 
disappear,  leaving  the  shafts  bare  from  end  to  end;  and  these  in 
the  tail  of  the  Paradisea  apoda  attain  a  length  of  thirty-four 
inches ;'^^  in  P.  Papuana  (fig.  47)  they  are  much  shorter  and  thin. 
Smaller  feathers  when  thus  denuded  appear  like  bristles,  as  on 
the  breast  of  the  turkey-cock.  As  any  fleeting  fashion  in  dress 
comes  to  be  admired  by  man,  so  with  birds  a  change  of  almost 
any  kind  in  the  structure  or  coloring  of  the  feathers  in  the  male 
appears  to  have  been  admired  by  the  female.  The  fact  of  the 
feathers  in  widely  distinct  groups,  having  been  modified  in  an 
analogous  manner,  no  doubt  depends  primarily  on  all  the  feathers 
having  nearly  the  same  structure  and  manner  of  development, 
and  consequently  tending  to  vary  in  the  same  manner.  We  often 
see  a  tendency  to  analogous  variability  in  the  plumage  of  our 
domestic  breeds  belonging  to  distinct  species.  Thus  top-knots 
have  appeared  in  several  species.  In  an  extinct  variety  of  the 
turkey,  the  top-knot  consisted  of  bare  quills  surmounted  with 
plumes  of  down,  so  that  they  somewhat  resembled  the  racket- 
shaped  feathers  above  described.  In  certain  breeds  of  the  pigeon 
and  fowl  the  feathers  are  plumose,  with  some  tendency  in  the 
shafts  to  be  naked.  In  the  Sebastopol  goose  the  scapular  feathers 
are  greatly  elongated,  curled,  or  even  spirally  twisted,  with  the 
margins  plumose.^^ 

In  regard  to  color  hardly  anything  need  here  be  said,  for 
every  one  knows  how  splendid  are  the  tints  of  many  birds,  and 
how  harmoniously  they  are  combined.  The  colors  are  often 
metallic  and  iridescent  Circular  spots  are  sometimes  surrounded 
by  one  or  more  differently  shaded  zones,  and  are  thus  converted 
into  ocelli.  Nor  need  much  be  said  on  the  wonderful  difference 
between  the  sexes  of  many  birds.  The  common  peacock  offers 
a  striking  instance.  Female  birds  of  paradise  are  obscurely  col- 
ored and  destitute  of  all  ornaments,  whilst  the  males  are  prob- 
ably the  most  highly  decorated  of  all  birds,  and  in  so  many  dif- 
ferent ways,  that  they  must  be  seen  to  be  appreciated.  The  elon- 
gated and  golden-orange  plumes  which  spring  from  beneath  the 

'0  'Proc.  Zoolog.  Soc'  1873,  p.  429. 

■^  Wallace,  in  'Annals  and  Mag-,  of  Nat.  Hist.'  vol.  xx.  1857,  p.  416; 
and  in  his  'Malay  Archipelag-o,  vol.  ii.  1869,  p.  390. 

'2  See  my  work  on  'The  Variation  of  Animals  and  Plants  under 
Domestication,'  vol.  1.  pp.  289,  293. 


DECORATION. 


381 


wings  of  the  Paradisea  apoda,  when  vertically  erected  and  made 
to  vibrate,  are  described  as  forming  a  sort  of  halo,  in  the  center 


Fig.  47.  Paradisea  Papuana  (T.  W.  Wood). 

of  v/hich  the  head  "looks  like  a  little  emerald  sun  with  its  rays 
"formed  by  the  two  plumes."^^    In  another  most  beautiful  species 


'8  Quoted  from  M.  de  Lafresnaye,  in  'Annals  and  Mag.  of  Nat.  Hist,' 
vol.  xiii.  1854,  p.  157;  see  also  Mr.  Wallace's  much  fuller  account  in 
vol.  XX.  1S57,  p.  412,  and  in  his  'Malay  Archipelago.' 


382  THE  DESCENT  OF  MAN. 

tlie  head  is  bald,  "and  of  a  rich  cobalt  blue,  crossed  by  several 
"lines  of  black  velvety  feathers."'* 

Male  humming-birds  (figs.  48  and  49)  almost  vie  with  birds  of 
paradise  in  their  beauty,  as  every  one  will  admit  who  has  seen 


Fig.  48.         Lophornls  ornatus,  male  and  female  (from  Brehm). 

Mr.  Gould's  splendid  volumes,  or  his  rich  collection.  It  is  very 
remarkable  in  how  many  different  ways  these  birds  are  orna- 
mented. Almost  every  part  of  their  plumage  has  been  taken 
advantage  of,  and  modified;  and  the  modifications  have  been 
carried,  as  Mr.  Gould  showed  me,  to  a  wonderful  extreme  in  some 
species   belonging  to   nearly   every  sub-group.     Such  cases   are 


71  Wallace,  'The  Malay  Archipelago,'  vol.  ii.  1869,  p.  405. 


DECORATION. 


383 


curiously  like  those  which  we  see  in  our  fancy  breeds,  reared 
by  man  for  the  sake  of  ornament:  certain  individuals  originally 
varied  in  one  character,  and  other  individuals  of  the  same  species 
in  other  characters;  and  these  have  been  seized  on  by  man 
and  much  augmented — as  shown  by  the  tail  of  the  fantail-pigeon. 


Fig.  49.       Spathura  underwoodi,  male  and  female  (from  Brehm). 

the  hood  of  the  jacobin,  the  beak  and  wattle  of  the  carrier,  and 
so  forth.  The  sole  difference  between  these  cases  is  that  in  the 
one,  the  result  is  due  to  man's  selection,  whilst  in  the  other,  as 
with  humming-birds,  birds  of  paradise,  &c.,  it  is  due  to  the  selec- 
tion by  the  females  of  the  more  beautiful  males. 

I  will  mention  only  one  other  bird,  remarkable  from  the  ex- 
treme contrast  in  color  between  the  sexes,  namely  the  famous 


384  THE  DESCENT  OF  MAN. 

bell-bird  (Chasmorhynclius  niveus)  of  S.  America,  the  note  of 
which  can  be  distinguished  at  the  distance  of  nearly  three  miles, 
and  astonishes  every  one  when  first  hearing  it.  The  male  is  pure 
white,  whilst  the  female  is  dusky-green;  and  white  is  a  very 
rare  color  in  terrestrial  species  of  moderate  size  and  inoffensive 
habits.  The  male,  also,  as  described  by  Waterton,  has  a  spiral 
tube,  nearly  three  inches  in  length,  which  rises  from  the  base 
of  the  beak.  It  is  jet-black,  dotted  over  with  minute  downy 
feathers.  This  tube  can  be  inflated  with  air,  through  a  communi- 
cation with  the  palate;  and  when  not  inflated  hangs  down  on 
one  side.  The  genus  consists  of  four  species,  the  males  of  which 
are  very  distinct,  whilst  the  females,  as  described  by  Mr.  Sclater 
in  a  very  interesting  paper,  closely  resemble  each  other,  thus 
offering  an  excellent  instance  of  the  common  rule  that  within 
the  same  group  the  males  differ  much  more  from  each  other 
than  do  the  females.  In  a  second  species  (C.  nudicollis)  the 
male  is  likewise  snow-white,  with  the  exception  of  a  large  space 
of  naked  skin  on  the  throat  and  round  the  eyes,  which  during 
the  breeding-season  is  of  a  fine  green  color.  In  a  third  species 
(C.  tricarunculatus)  the  head  and  neck  alone  of  the  male  are 
white,  the  rest  of  the  body  being  chestnut-brown,  and  the  male 
of  this  species  is  provided  with  three  filamentous  projections 
half  as  long  as  the  body — one  rising  from  the  base  of  the  beak, 
and  the  two  others  from  the  corners  of  the  mouth." 

The  colored  plumage  and  certain  other  ornaments  of  the  adult 
males  are  either  retained  for  life,  or  are  periodically  renewed 
during  the  summer  and  breeding-season.  At  this  same  season 
the  beak  and  naked  skin  about  the  head  frequently  change  color, 
as  with  some  herons,  ibises,  gulls,  one  of  the  bell-birds  just  no- 
ticed, &c.  In  the  white  ibis,  the  cheeks,  the  inflatable 
skin  of  the  throat,  and  the  basal  portion  of  the  beak  then  be- 
come crimson.'®  In  one  of  the  rails,  Gallicrex  cristatus,  a  large  red 
caruncle  is  developed  during  this  period  on  the  head  of  the  male. 
So  it  is  with  a  thin  horny  crest  on  the  beak  of  one  of  the  pelicans, 
P.  erythrorhynchus;  for  after  the  breeding-season,  these  horny 
crests  are  shed,  like  horns  from  the  heads  of  stags,  and  the  shore 
of  an  island  in  a  lake  in  Nevada  was  found  covered  with  these 
curious  exuviae." 

Changes  of  color  in  the  plumage  according  to  the  season  de- 
pend, firstly  on  a  double  annual  moult,  secondly  on  an  actual 
change  of  color  in  the  feathers  themselves,  and  thirdly  on  their 

■^s  Mr.  Sclater,  'Intellectual  Observer,'  Jan.  1867.  'Waterton' s  Wan- 
derings,' p.  118.  See  also  Mr.  Salvin's  interesting  paper,  with  a  plate, 
in  the  'Ibis,'  1865,  p.  90. 

^e  'Land  and  Water,'  1867,  p.  394. 

"  Mr.  D.  G.  ElUot,  in  'Proc.  Zool.  Soc'  1869,  p.  589. 


DOUBLE  ANNUAL  MOULT.  385 

dull-colored  margins  being  periodically  shed,  or  on  these  three 
processes  more  or  less  combined.  The  shedding  of  the  deciduary 
margins  may  be  compared  with  the  shedding  of  their  down  by 
very  young  birds;  for  the  down  in  most  cases  arises  from  the 
summits  of  the  first  true  feathers.'^^ 

With  respect  to  the  birds  which  annually  undergo  a  double 
moult,  there  are,  firstly,  some  kinds,  for  instance  snipes,  swallow- 
plovers  (Glareolse),  and  curlews,  in  which  the  two  sexes  resemble 
each  other,  and  do  not  change  color  at  any  season.  I  do  not 
know  whether  the  winter  plumage  is  thicker  and  warmer  than 
the  summer  plumage,  but  warmth  seems  the  most  probable  end 
attained  of  a  double  moult,  where  there  is  no  change  of  color. 
Secondly,  there  are  birds,  for  instance,  certain  species  of  Totanus 
and  other  Grallatores,  the  sexes  of  which  resemble  each  other, 
but  in  which  the  summer  and  winter  plumage  differ  slightly  in 
color.  The  difference,  however,  in  these  cases  is  so  small  that 
it  can  hardly  be  an  advantage  to  them;  and  it  may,  perhaps, 
be  attributed  to  the  direct  action  of  the  different  conditions  to 
which  the  birds  are  exposed  during  the  two  seasons.  Thirdly, 
there  are  many  other  birds  the  sexes  of  which  are  alike,  but 
which  are  widely  different  in  their  summer  and  winter  plumage. 
Fourthly,  there  are  birds,  the  sexes  of  which  differ  from  each 
other  in  color;  but  the  females,  though  moulting  twice,  retain 
the  same  colors  throughout  the  year,  whilst  the  males  undergo  a 
change  of  color,  sometimes  a  great  one,  as  with  certain  bustards. 
Fifthly  and  lastly,  there  are  birds  the  sexes  of  which  differ  from 
each  other  in  both  their  summer  and  winter  plumage;  but  the 
male  undergoes  a  greater  amount  of  change  at  each  recurrent 
season  than  the  female — of  which  the  ruff  (Machetes  pugnax) 
offers  a  good  instance. 

With  respect  to  the  cause  or  purpose  of  the  differences  in 
color  between  the  summer  and  winter  plumage,  this  may  in 
some  instances,  as  with  the  ptarmigan,^^  serve  during  both  sea- 
sons as  a  protection.  When  the  difference  between  the  two 
plumages  is  slight,  it  may  perhaps  be  attributed,  as  already  re- 
marked, to  the  direct  action  of  the  conditions  of  life.  But  with 
many  birds  there  can  hardly  be  a  doubt  that  the  summer  plumage 
is  ornamental,  even  when  both  sexes  are  alike.  We  may  con- 
clude that  this  is  the  case  with  many  herons,  egrets,  &c.,  for 

■^8  Nitzsch's  'Pterylography,'  edited  by  P.  L,  Sclater.  Ray  Soc.  1867, 
p.  U. 

''^  The  brown  mottled  summer  plumage  of  the  ptarmigan  is  of  as 
much  importance  to  it,  as  a  protection,  as  the  white  winter  plumage; 
for  in  Scandinavia,  during  the  spring,  when  the  snow  has  disappeared, 
this  bird  is  known  to  suffer  greatly  from  birds  of  prey,  before  it  has 
acquired  its  summer  dress:  see  Wilhelm  von  "Wright,  in  Lloyd,  'Game 
Birds  of  Sweden,  1867,  p.  125. 
28 


386  THE  DESCENT  OF  MAN. 

they  acquire  their  beautiful  plumes  only  during  the  breeding- 
season.  Moreover,  such  plumes,  top-knots,  &c.,  though  possessed 
by  both  sexes,  are  occasionally  a  little  more  developed  in  the 
male  than  in  the  female;  and  they  resemble  the  plumes  and 
ornaments  possessed  by  the  males  alone  of  other  birds.  It  is  also 
known  that  confinement,  by  affecting  the  reproductive  system  of 
male  birds,  frequently  checks  the  development  of  their  secondary 
sexual  characters,  but  has  no  immediate  influence  on  any  other 
characters;  and  I  am  informed  by  Mr.  Bartlett  that  eight  or 
nine  specimens  of  the  Knot  (Tringa  canutus)  retained  their  un- 
adorned winter  plumage  in  the  Zoological  Gardens  throughout 
the  year,  from  which  fact  we  may  infer  that  the  summer  plumage 
though  common  to  both  sexes  partakes  of  the  nature  of  the  ex- 
clusively masculine  plumage  of  many  other  birds.^*' 

From  the  foregoing  facts,  more  especially  from  neither  sex  of 
certain  birds  changing  color  during  either  annual  moult,  or 
changing  so  slightly  that  the  change  can  hardly  be  of  any  service 
to  them,  and  from  the  females  of  other  species  moulting  twice 
yet  retaining  the  same  colors  throughout  the  year,  we  may 
conclude  that  the  habit  of  annually  moulting  twice  has  not  been 
acquired  in  order  that  the  male  should  assume  an  ornamental 
character  during  the  breeding-season;  but  that  the  double  moult, 
having  been  originally  acquired  for  some  distinct  purpose,  has 
subsequently  been  taken  advantage  of  in  certain  cases  for  gain- 
ing a  nuptial  plumage. 

It  appears  at  first  sight  a  surprising  circumstance  that  some 
closely-allied  species  should  regularly  undergo  a  double  annual 
moult,  and  others  only  a  single  one.  The  ptarmigan,  for  instance, 
moults  twice  or  even  thrice  in  the  year,  and  the  black-cock  only 
once:  some  of  the  splendidly  colored  honey-suckers  (Nectariniae) 
of  India  and  some  sub-genera  of  obscurely  colored  pipits  (Anthus) 
have  a  double,  whilst  others  have  only  a  single  annual  moult.*^ 
But  the  gradations  in  the  manner  of  moulting,  which  are  known 
to  occur  with  various  birds,  show  us  how  species,  or  whole  groups, 
might  have  originally  acquired  their  double  annual  moult,  or 
having  once  gained  the  habit,  have  again  lost  it.  With  certain 
bustards  and  plovers  the  vernal  moult  is  far  from  complete,  some 
feathers  being  renewed,  and  some  changed  in  color.     There  is 


80  In  regai'd  to  the  previous  statements  on  moulting,  see,  on  snipes, 
&c.,  Macgillivray,  'JEIist.  Brit.  Birds/  vol.  iv.  p.  371;  on  Glareolae,  cur- 
lews, and  bustards,  Jerdon,  'Birds  of  India,'  vol.  iii.  pp.  615,  630,  683; 
on  Totanus,  ibid.  p.  700;  on  the  plumes  of  herons,  ibid.  p.  738,  and 
Macgillivray,  vol.  iv.  pp.  435  and  444,  and  Mr.  Stafford  Allen,  in  the 
'Ibis,'  vol.  V.  1863,  p.  S3. 

81  On  the  moulting-  of  the  ptarmigan,  see  Gould's  'Birds  of  Great 
Britain.'  On  the  honey-suckers,  Jerdon,  'Birds  of  India,'  vol.  i.  pp. 
359,  365,  369.    On  the  moulting  of  Anthus,  see  Blyth,  in  'Ibis,'  1867,  p.  32. 


DOUBLE  ANNUAL  MOULT.  387 

also  reason  to  believe  that  with  certain  bustards  and  rail-like 
birds,  which  properly  undergo  a  double  moult,  some  of  the  older 
males  retain  their  nuptial  plumage  throughout  the  year.  A  few 
highly  modified  feathers  may  merely  be  added  during  the  spring 
to  the  plumage,  as  occurs  with  the  disc-formed  tail-feathers  of 
certain  drongos  (Bhringa)  in  India,  and  with  the  elongated 
feathers  on  the  back,  neck,  and  crest  of  certain  herons.  By  such 
steps  as  these,  the  vernal  moult  might  be  rendered  more  and 
more  complete,  until  a  perfect  double  moult  was  acquired.  Some 
of  the  birds  of  paradise  retain  their  nuptial  feathers  throughout 
the  year,  and  thus  have  only  a  single  moult;  others  cast  them  di- 
rectly after  the  breeding-season,  and  thus  have  a  double  moult; 
and  others  again  cast  them  at  this  season  during  the  first  year, 
but  not  afterwards;  so  that  these  latter  species  are  intermediate 
in  their  manner  of  moulting.  There  is  also  a  great  difference 
with  many  birds  in  the  length  of  time  during  which  the  two 
annual  plumages  are  retained;  so  that  the  one  might  come  to  be 
retained  for  the  whole  year,  and  the  other  completely  lost.  Thus 
in  the  spring  Machetes  pugnax  retains  his  ruff  for  barely  two 
months.  In  Natal  the  male  widow-bird  (Chera  progne)  acquires 
his  fine  plumage  and  long  tail-feathers  in  December  or  January, 
and  loses  them  in  March;  so  that  they  are  retained  only  for 
about  three  months.  Most  species,  which  undergo  a  double  moult, 
keep  their  ornamental  feathers  for  about  six  months.  The  male, 
however,  of  the  wild  Gallus  bankiva  retains  his  neck-hackles 
for  nine  or  ten  months;  and  when  these  are  cast  off,  the  under- 
lying black  feathers  on  the  neck  are  fully  exposed  to  view.  But 
with  the  domesticated  descendant  of  this  species,  the  neck-hackles 
of  the  male  are  immediately  replaced  by  new  ones;  so  that  we 
here  see,  as  to  part  of  the  plumage,  a  double  moult  changed  under 
domestication  into  a  single  moult.®- 

The  common  drake  (Anas  boschas)  after  the  breeding-season 
is  well  known  to  lose  his  male  plumage  for  a  period  of  three 
months,  during  which  time  he  assumes  that  of  the  female.  The 
male  pintail-duck  (Anas  acuta)  loses  his  plumage  for  the  shorter 
period  of  six  weeks  or  two  months;  and  Montagu  remarks  that 
"this  double  moult  within  so  short  a  time  is  a  most  extraordinary 

82  For  the  foregoing  statements  in  regard  to  partial  moults,  and  on 
old  males  retaining  their  nuptial  plumage,  see  Jerdon,  on  bustards 
and  plovers,  in  'Birds  of  India,'  vol.  iii.  pp.  617,  637,  709,  711.  Also  Blyth 
in  'Land  and  Water,'  1867,  p.  84.  On  the  moulting  of  Paradisea,  see  an 
interesting  article  by  Dr.  W.  Marshall,  'Archives  Neerlandaises,'  torn. 
vi.  1871.  On  the  Vidua,  'Ibis,'  vol.  iii.  1861,  p.  133.  On  the  Drongo-shrlkes, 
Jerdon,  ibid,  vol  i.  p.  435.  On  the  vernal  moult  of  the  Herodias  bubul- 
cus,  Mr.  S.  S.  Allen,  in  'Ibis,'  1863,  p.  33.  On  Gallus  bankiva,  Blyth,  in 
'Annals  and  Mag.  of  Nat.  Hist.'  vol.  i.  1848,  p.  455;  see,  also,  on  this 
subject,  my  'Variation  of  Animals  under  Domestication,'  vol.  i.  p.  236. 


388  THE   DESCENT  OF  MAN. 

"circumstance,  that  seems  to  bid  defiance  to  all  human  reason- 
"ing."  But  the  believer  in  the  gradual  modification  of  species  will 
be  far  from  feeling  surprise  at  finding  gradations  of  all  kinds. 
If  the  male  pintail  were  to  acquire  his  new  plumage  within  a 
still  shorter  period,  the  new  male  feathers  would  almost  neces- 
sarily be  mingled  with  the  old,  and  both  with  some  proper  to  the 
female;  and  this  apparently  is  the  case  with  the  male  of  a  not 
distantly-allied  bird,  namely  the  Merganser  serrator,  for  the 
males  are  said  to  "undergo  a  change  of  plumage,  which  assimi- 
"lates  them  in  some  measure  to  the  female."  By  a  little  further 
acceleration  in  the  process,  the  double  moult  would  be  completely 
iost.^ 

Some  male  birds,  as  before  stated,  become  more  brightly  col- 
ored in  the  spring,  not  by  a  vernal  moult,  but  either  by  an  actual 
change  of  color  in  the  feathers,  or  by  their  obscurely-colored 
deciduary  margins  being  shed.  Changes  of  color  thus  caused  may 
last  for  a  longer  or  shorter  time.  In  the  Pelecanus  onocrotalus 
a  beautiful  rosy  tint,  with  lemon-colored  marks  on  the  breast, 
overspreads  the  whole  plumage  in  the  spring;  but  these  tintfe,  as 
Mr.  Sclater  states,  "do  not  last  long,  disappearing  generally  in 
"about  six  weeks  or  two  months  after  they  have  been  attained." 
Certain  finches  shed  the  margins  of  their  feathers  in  the  spring, 
and  then  become  brighter  colored,  while  other  finches  undergo 
no  such  change.  Thus  the  Fringilla  tristis  of  the  United  States 
(as  well  as  many  other  American  species)  exhibits  its  bright 
colors  only  when  the  winter  is  past,  whilst  our  goldfinch,  which 
exactly  represents  this  bird  in  habits,  and  our  siskin,  which  rep- 
resents it  still  more  closely  in  structure,  undergo  no  such  annual 
change.  But  a  difference  of  this  kind  in  the  plumage  of  allied 
species  is  not  surprising,  for  with  the  common  linnet,  which 
belongs  to  the  same  family,  the  crimson  forehead  and  breast  are 
displayed  only  during  the  summer  in  England,  whilst  in  Madeira 
these  colors  are  retained  throughout  the  year.^* 

Display  by  Male  Birds  of  their  Plumage. — Ornaments  of  all 
kinds,  whether  permanently  or  temporarily  gained,  are  sedulously 
displayed  by  the  males,  and  apparently  serve  to  excite,  attract, 
or  fascinate  the  females.  But  the  males  will  sometimes  display 
their  ornaments,  when  not  in  the  presence  of  the  females,   as 

83  See  Macg-illivray,  'Hist.  British  Birds'  (vol.  v.  pp.  34,  70,  and  223), 
on  the  moulting  of  the  Anatidae,  with  quotations  from  Waterton  and 
Montagu.    Also  Yarrell,  'Hist,  of  British  Birds,'  vol.  iii.  p.  243. 

84  On  the  pelican,  see  Sclater,  in  'Proc.  Zool.  Soc'  1868,  p.  265.  On 
the  American  finches,  see  Audubon,  'Ornith.  Biography,'  vol.  i.  pp. 
174,  221,  and  Jerdon,  'Birds  of  India,'  vol.  ii.  p.  383.  On  the  Fringilla 
cannabina  of  Madeira,  Mr.  E.  Vernon  Harcourt,  'Ibis,'  vol.  v.,  1863, 
p.  230. 


DISPLAY  BY  THE  MALE. 


389 


occasionally  occurs  with  grouse  at  their  balz-places,  and  as  may- 
be noticed  with  the  peacock;  this  latter  bird,  however,  evidently 
wishes  for  a  spectator  of  some  kind,  and,  as  I  have  often  seen, 
will  show  off  his  finery  before  poultry,  or  even  pigs.^  AH  natu- 
ralists who  have  closely  attended  to  the  habits  of  birds,  whether 
in  a  state  of  nature  or  under  confinement,  are  unanimously  of 
opinion  that  the  males  take  delight  in  displaying  their  beauty. 
Audubon  frequently  speaks  of  the  male  as  endeavoring  in  various 
ways  to   charm  the  female.     Mr.   Gould,  after  describing  some 


Fig-.  50. 


Rupicola  crocea,  male  (T.  W.  Wood). 


peculiarities  in  a  male  humming-bird,  says  he  has  no  doubt  that 
it  has  the  power  of  displaying  them  to  the  greatest  advantage 
before  the  female.  Dr.  Jerdon^^  insists  that  the  beautiful  plumage 
of  the  male  serves  "to  fascinate  and  attract  the  female."  Mr. 
Bartlett,  at  the  Zoological  Gardens,  expressed  himself  to  me  in 
the  strongest  terms  to  the  same  effect. 


85  See  also  'Ornamental  Poultry,'  by  Rev.  E.  S.  Dixon,  1848,  p.  8. 

86  'Birds  of  India,'  ^jitroduct.  vol.  i.  p.  xxiv.;  on  the  peacock,  vol. 
iii.  p.  507.  See  Gould's  'Introduction  to  the  Trochilidae,'  1861,  pp.  15 
and  HI. 


26 


390  THE  DESCENT  OF  MAN. 

It  must  be  a  grand  sight  in  the  forests  of  India  "to  come  sud- 
"denly  on  twenty  or  thirty  pea-fowl,  the  males  displaying  their 
"gorgeous  trains,  and  strutting  about  in  all  the  pomp  of  pride 
"before  the  gratified  females."  The  wild  turkey-cock  erects  his 
glittering  plumage,  expands  his  finely-zoned  tail  and  barred 
wing-feathers,  and  altogether,  with  his  crimson  and  blue  wattles, 
makes  a  superb,  though  to  our  eyes,  grotesque  appearance.  Simi- 
lar facts  have  already  been  given  with  respect  to  grouse  of  va- 
rious kinds.  Turning  to  another  Order.  The  male  Rupicola  cro- 
cea  (fig.  50)  is  one  of  the  most  beautiful  birds  in  the  world,  being 
of  a  splendid  orange,  with  some  of  the  feathers  curiously  trun- 
cated and  plumose.  The  female  is  brownish-green,  shaded  with 
red,  and  has  a  much  smaller  crest.  Sir  R.  Schomburgk  has  de- 
scribed their  courtship;  he  found  one  of  their  meeting-places 
where  ten  males  and  two  females  v/ere  present.  The  space  was 
from  four  to  five  feet  in  diameter,  and  appeared  to  have  been 
cleared  of  every  blade  of  grass  and  smoothed  as  if  by  human 
hands.  A  male  "was  capering,  to  the  apparent  delight  of  several 
"others.  Now  spreading  its  wings,  throwing  up  its  head,  or  open- 
"ing  its  tail  like  a  fan;  now  strutting  about  with  a  hopping  gait 
"until  tired,  when  it  gabbled  some  kind  of  note,  and  was  relieved 
"by  another.  Thus  three  of  them  successively  took  the  field, 
"and  then,  with  self-approbation,  withdrew  to  rest."  The  In- 
dians, in  order  to  obtain  their  skins,  wait  at  one  of  the  meeting- 
places  till  the  birds  are  eagerly  engaged  in  dancing,  and  then 
are  able  to  kill  with  their  poisoned  arrows  four  or  five  males, 
one  after  the  other. ^^  With  birds  of  paradise  a  dozen  or  more 
full-plumaged  males  congregate  in  a  tree  to  hold  a  dancing-party, 
as  it  is  called  by  the  natives:  and  here  they  fly  about,  raise  their 
wings,  elevate  their  exquisite  plumes,  and  make  them  vibrate, 
and  the  whole  tree  seems,  as  Mr.  Wallace  remarks,  to  be  filled 
with  waving  plumes.  When  thus  engaged,  they  become  so  ab- 
sorbed that  a  skillful  archer  may  shoot  nearly  the  whole  party. 
These  birds,  when  kept  in  confinement  in  the  Malay  Archipelago, 
are  said  to  take  much  care  in  keeping  their  feathers  clean;  often 
spreading  them  out,  examining  them,  and  removing  every  speck 
of  dirt.  One  observer,  who  kept  several  pairs  alive,  did  not 
doubt  that  the  display  of  the  male  was  intended  to  please  tiie 
female.^ 

The  Gold  and  Amherst  pheasants  during  their  courtship  not 
only  expand  and  raise  their  splendid  frills,  but  twist  them,  as  I 
have  myself  seen,  obliquely  towards  the  female  on  whichever  side 


87  'Journal  of  R.  Geograph.  Soc'  vol.  x.  1840,  p.  236. 

88  "Annals  and  Mag.  of  Nat.  Hist.'  vol.  xiii.  1854,  p.  157;  also  Wallace, 
ibid.  vol.  XX.  1857,  p.  412,  and  'The  Malay  Archipelago,'  vol.  ii.  1869,  p. 
252.   Also  Dr.  Bennett,  as  quoted  by  Brehm,  'Thierleben,*  B.  iii.  s.  326. 


DISPLAY  BY  THE  MALE. 


391 


she  maj-  be  standing,  obviously  in  order  that  a  large  surface  may 
be  displayed  before  her.^®  They  likewise  turn  their  beautiful 
tails  and  tail-coverts  a  little  towards  the  same  side.    Mr.  Bartlett 


Fig.  51 


Polyplectron  chinquis,  male  (T.  W.  Wood). 


89  Mr.  T.  W.  Wood  has  given  ('The  Student,'  April  1870.  p.  115)  a  full 
account  of  this  manner  of  display,  by  the  Gold  pheasant  and  by  the 
Japanese  pheasant,  Ph.  versicolor:  and  he  calls  it  the  lateral  or  one- 
sided display. 


392  THE  DESCENT  OF  MAN. 

has  observed  a  male  Polyplectron  (fig.  51)  in  the  act  of  court- 
ship, and  has  shown  me  a  specimen  stuffed  in  the  attitude  then 
assumed.  The  tail  and  wing-feathers  of  this  bird  are  ornamented 
with  beautiful  ocelli,  like  those  on  the  peacock's  train.  Now 
when  the  peacock  displays  himself,  he  expands  and  erects  his 
tail  transversely  to  his  body,  for  he  stands  in  front  of  the  female, 
and  has  to  show  off,  at  the  same  time,  his  rich  blue  throat  and 
breast.  But  the  breast  of  the  Polyplectron  is  obscurely  colored, 
and  the  ocelli  are  not  confined  to  the  tail-feathers.  Consequently 
the  Polyplectron  does  not  stand  in  front  of  the  female;  but  he 
erects  and  expands  his  tail-feathers  a  little  obliquely,  lowering 
the  expanded  wing  on  the  same  side,  and  raising  that  on  the 
opposite  side.  In  this  attitude  the  ocelli  over  the  whole  body 
are  exposed  at  the  same  time  before  the  eyes  of  the  admiring 
female  in  one  grand  bespangled  expanse.  To  whichever  side  she 
may  turn,  the  expanded  wings  and  the  obliquely-held  tail  are 
turned  towards  her.  The  male  Tragopan  pheasant  acts  in  nearly 
the  same  manner,  for  he  raises  the  feathers  of  the  body,  though 
not  the  wing  itself,  on  the  side  which  is  opposite  to  the  female, 
and  which  would  otherwise  be  concealed,  so  that  nearly  all  the 
beautifully  spotted  feathers  are  exhibited  at  the  same  time. 

The  Argus  pheasant  affords  a  much  more  remarkable  case.  The 
immensely  developed  secondary  wing-feathers  are  confined  to  the 
male;  and  each  is  ornamented  with  a  row  of  from  twenty  to 
twenty-three  ocelli,  above  an  inch  in  diameter.  These  feathers 
are  also  elegantly  marked  with  oblique  stripes  and  rows  of  spots 
of  a  dark  color,  like  those  on  the  skin  of  a  tiger  and  leopard 
combined.  These  beautiful  ornaments  are  hidden  until  the  male 
shows  himself  off  before  the  female.  He  then  erects  his  tail,  and 
expands  his  wing-feathers  into  a  great,  almost  upright,  circular 
fan  or  shield,  which  is  carried  in  front  of  the  body.  The  neck 
and  head  are  held  on  one  side,  so  that  they  are  concealed  by  the 
fan;  but  the  bird  in  order  to  see  the  female,  before  wliom  he  is 
displaying  himself,  sometimes  pushes  his  head  between  two  of 
the  long  wing-feathers  (as  Mr.  Bartlett  has  seen),  and  then  pre- 
sents a  grotesque  appearance.  This  must  be  a  frequent  habit 
with  the  bird  in  a  state  of  nature,  for  Mr.  Bartlett  and  his  son  on 
examining  some  perfect  skins  sent  from  the  East,  found  a  place 
between  two  of  the  feathers,  which  was  much  frayed,  as  if  the 
head  had  here  frequently  been  pushed  through.  Mr.  Wood  thinks 
that  the  male  can  also  peep  at  the  female  on  one  side,  beyond 
the  margin  of  the  fan. 

The  ocelli  on  the  wing-feathers  are  wonderful  objects;  for  they 
are  so  shaded  that,  as  the  Duke  of  Argyll  remarks,^"  they  stand 

^  'The  Reign  of  Law,'  1867,  p.  203. 


DISPLAY  BY  THE  MALE. 


393 


out  like  balls  lying  loosely  within  sockets.  When  I  looked  at 
the  specimen  in  the  British  Museum,  which  is  mounted  with  the 
wings  expanded  and  trailing  downwards,  I  was  however  greatly 


Fig.  52.   Side  view  of  male  Argus  pheasant,  whilst  displaying  before  the 
female.    Observed  and  sketched  from  nature  by  Mr,  T.  W.  Wood. 

disappointed,  for  the  ocelli  appeared  flat,  or  even  concave.     But 
Mr.   Gould  soon  made  the  case  clear  to   me,  for  he  held  the 


394  THE  DESCENT  OF  MAN. 

feathers  erect,  in  the  position  in  which  they  would  naturally  be 
displayed,  and  now,  from  the  light  shining  on  them  from  above, 
each  ocellus  at  once  resembled  the  ornament  called  a  ball  and 
socket.  These  feathers  have  been  shown  to  several  artists,  and 
all  have  expressed  their  admiration  at  the  perfect  shading.  It 
may  well  be  asked,  could  such  artistically  shaded  ornaments  have 
been  formed  by  means  of  sexual  selection?  But  it  will  be  con- 
venient to  defer  giving  an  answer  to  this  question,  until  we  treat 
in  the  next  chapter  of  the  principle  of  gradation. 

The  foregoing  remarks  relate  to  the  secondary  wing-feathers, 
but  the  primary  wing-feathers,  which  in  most  gallinaceous  birds 
are  uniformly  colored,  are  in  the  Argus  pheasant  equally  won- 
derful. They  are  of  a  soft  brown  tint  with  numerous  dark  spots, 
each  of  which  consists  of  two  or  three  black  dots  with  a  surround- 
ing dark  zone.  But  the  chief  ornament  is  a  space  parallel  to  the 
dark-blue  shaft,  which  in  outline  forms  a  perfect  second  feather 
lying  within  the  true  feather.  This  inner  part  is  colored  of  a 
lighter  chestnut,  and  is  thickly  dotted  with  minute  white  points. 

I  have  shown  this  feather  to  several  persons,  and  many  have 
admired  it  even  more  than  the  ball  and  socket  feathers,  and 
have  declared  that  it  was  more  like  a  work  of  art  than  of  nature. 
Now  these  feathers  are  quite  hidden  on  all  ordinary  occasions, 
but  are  fully  displayed,  together  with  the  long  secondary  feathers, 
when  they  are  all  expanded  together  so  as  to  form  the  great  fan  or 
shield. 

The  case  of  the  male  Argus  pheasant  is  eminently  interesting, 
because  it  affords  good  evidence  that  the  most  refined  beauty  may 
serve  as  a  sexual  charm,  and  for  no  other  purpose.  We  must  con- 
clude that  this  is  the  case,  as  the  secondary  and  primary  wing- 
feathers  are  not  at  all  displayed,  and  the  ball  and  socket  orna- 
ments are  not  exhibited  in  full  perfection,  until  the  male  assumes 
the  attitude  of  courtship.  The  Argus  pheasant  does  not  possess 
brilliant  colors,  so  that  his  success  in  love  appears  to  depend  on 
the  great  size  of  his  plumes,  and  on  the  elaboration  of  the  most 
elegant  patterns.  Many  will  declare  that  it  is  utterly  incredible 
that  a  female  bird  should  be  able  to  appreciate  fine  shading  and 
exquisite  patterns.  It  is  undoubtedly  a  marvelous  fact  that  she 
should  possess  this  almost  human  degree  of  taste.  He  who  thinks 
that  he  can  safely  gauge  the  discrimination  and  taste  of  the 
lower  animals  may  deny  that  the  female  Argus  pheasant  can  ap- 
preciate such  refined  beauty;  but  he  will  then  be  compelled  to  ad- 
mit that  the  extraordinary  attitudes  assumed  by  the  male  during 
the  act  of  courtship,  by  which  the  wonderful  beauty  of  his  plum- 
age is  fully  displayed,  are  purposeless;  and  this  is  a  conclusion 
which  I  for  one  will  never  admit. 

Although  so  many  pheasants  and  allied  gallinaceous  birds  care- 
fully display  their  plumage  before  the  females,  it  is  remarkable, 


DISPLAY  BY  THE  MALE.  395 

as  Mr.  Bartlett  informs  me,  that  this  is  not  the  case  with  the 
dull-colored  Eared  and  Cheer  pheasants  (Crossoptilon  auritum 
and  Phasianus  wallichii);  so  that  these  birds  seem  conscious  that 
they  have  little  beauty  to  display.  Mr.  Bartlett  has  never  seen 
the  males  of  either  of  these  species  fighting  together,  though 
he  has  not  had  such  good  opportunities  for  observing  the  Cheer 
as  the  Eared  pheasant.  Mr.  Jenner  Weir,  also,  finds  that  all 
male  birds  with  rich  or  strongly-characterized  plumage  are  more 
quarrelsome  than  the  dull-colored  species  belonging  to  the  same 
groups.  The  goldfinch,  for  instance,  is  far  more  pugnacious  than 
the  linnet,  and  the  blackbird  than  the  thrush.  Those  birds  which 
undergo  a  seasonal  change  of  plumage  likewise  become  much 
more  pugnacious  at  the  period  when  they  are  most  gayly  orna- 
mented. No  doubt  the  males  of  some  obscurely-colored  birds 
fight  desperately  together,  but  it  appears  that  when  sexual  selec- 
tion has  been  highly  influential,  and  has  given  bright  colors  to  the 
males  of  any  species,  it  has  also  very  often  given  a  strong  ten- 
dency to  pugnacity.  We  shall  meet  with  nearly  analogous  cases 
when  we  treat  of  mammals.  On  the  other  hand,  with  birds  the 
power  of  song  and  brilliant  colors  have  rarely  been  both  acquired 
by  the  males  of  the  same  species;  but  in  this  case,  the  advantage 
gained  would  have  been  the  same,  namely,  success  in  charming 
the  female.  Nevertheless  it  must  be  owned  that  the  males  of 
several  brilliantly  colored  birds  have  had  their  feathers  specially 
modified  for  the  sake  of  producing  instrumental  music,  though 
the  beauty  of  this  cannot  be  compared,  at  least  according  to  our 
taste,  with  that  of  the  vocal  music  of  many  songsters. 

We  will  now  turn  to  male  birds  which  are  not  ornamented  in 
any  high  degree,  but  which  nevertheless  display  during  their 
courtship  whatever  attractions  they  may  possess.  These  cases 
are  in  some  respects  more  curious  than  the  foregoing,  and  have 
been  but  little  noticed.  I  owe  the  following  facts  to  Mr.  Weir, 
who  has  long  kept  confined  birds  of  many  kinds,  including  all  the 
British  Fringillidae  and  Emberizida3.  The  facts  have  been  selected 
from  a  large  body  of  valuable  notes  kindly  sent  me  by  him.  The 
bullfinch  makes  his  advances  in  front  of  the  female,  and  then 
puffs  out  his  breast,  so  that  many  more  of  the  crimson  feathers 
are  seen  at  once  than  otherwise  would  be  the  case.  At  the 
same  time  he  twists  and  bows  his  black  tail  from  side  to  side  in 
a  ludicrous  manner.  The  male  chaffinch  also  stands  in  front  of 
the  female,  thus  showing  his  red  breast  and  "blue  bell,"  as  the 
fanciers  call  his  head;  the  wings  at  the  same  time  being  slightly 
expanded,  with  the  pure  white  bands  on  the  shoulders  thus  ren- 
dered conspicuous.  The  common  linnet  distends  his  rosy  breast, 
slightly  expands  his  brown  wings  and  tail,  so  as  to  make  the 
best  of  them  by  exhibiting  their  white  edgings.  We  must,  how- 
ever, be  cautious  in  concluding  that  the  wings  are  spread  out 


S96  THE  DESCENT  OF  MAN. 

solely  for  display,  as  some  birds  do  so  whose  wings  are  not  beau- 
tiful. This  is  the  case  with  the  domestic  cock,  but  it  is  always 
the  wing  on  the  side  opposite  to  the  female  which  is  expanded, 
and  at  the  same  time  scraped  on  the  ground.  The  male  goldfinch 
behaves  differently  from  all  other  finches:  his  wings  are  beau- 
tiful, the  shoulders  being  black,  with  the  dark-tipped  wing-feath- 
ers spotted  with  white  and  edged  with  golden  yellow.  When  he 
courts  the  female,  he  sways  his  body  from  side  to  side,  and 
quickly  turns  his  slightly  expanded  wings  first  to  one  side,  then 
to  the  other,  with  a  golden  flashing  effect.  Mr.  Weir  informs  me 
that  no  other  British  finch  turns  thus  from  side  to  side  during 
his  courtship,  not  even  the  closely-allied  male  siskin,  for  he 
would  not  thus  add  to  his  beauty. 

Most  of  the  British  Buntings  are  plain  colored  birds;  but  in  the 
spring  the  feathers  on  the  head  of  the  male  reed-bunting  (Em- 
beriza  schoeniculus)  acquire  a  fine  black  color  by  the  abrasion  of 
the  dusky  tips;  and  these  are  erected  during  the  act  of  courtship. 
Mr.  Weir  has  kept  two  species  of  Amadina  from  Australia:  the 
A.  castanotis  is  a  very  small  and  chastely  colored  finch,  with  a 
dark  tail,  white  rump,  and  jet-black  upper  tail-coverts,  each  of 
the  latter  being  marked  with  three  large  conspicuous  oval  spots 
of  white.''^  This  species,  when  courting  the  female,  slightly 
spreads  out  and  vibrates  these  parti-colored  tail-coverts  in  a  very 
peculiar  manner.  The  male  Amadina  Lathami  behaves  very  dif- 
ferently, exhibiting  before  the  female  his  brilliantly  spotted 
breast,  scarlet  rump,  and  scarlet  upper  tail-coverts.  I  may  here 
add  from  Dr.  Jerdon  that  the  Indian  bulbul  (Pycnonotus  hsemor- 
rhous)  has  its  under  tail-coverts  of  a  crimson  color,  and  these,  it 
might  be  thought,  could  never  be  well  exhibited;  but  the  bird 
"when  excited  often  spreads  them  out  laterally,  so  that  they  can  be 
"seen  even  from  above."^^  The  crimson  under  tail-coverts  of  some 
other  birds,  as  with  one  of  the  woodpeckers,  Picus  major,  can  be 
seen  without  any  such  display.  The  common  pigeon  has  Iridescent 
feathers  on  the  breast,  and  every  one  must  have  seen  how  the  male 
infiates  his  breast,  whilst  courting  the  female,  thus  showing  them 
off  to  the  best  advantage.  One  of  the  beautiful  bronze-winged  pig- 
eons of  Australia  (Ocyphaps  lophotes)  behaves,  as  described  to  me 
by  Mr.  Weir,  very  differently:  the  male,  whilst  standing  before 
the  female,  lowers  his  head  almost  to  the  ground,  spreads  out 
and  raises  his  tail,  and  half  expands  his  wings.  He  then  alter- 
nately and  slowly  raises  and  depresses  his  body,  so  that  the 
iridescent  metallic  feathers  are  all  seen  at  once,  and  glitter  in 
the  sun. 

»i  For  the  description  of  these  birds,  see  Gould's  'Handbook  to  the 
Birds  of  Australia,'  vol.  i.  1865,  p.  417. 
»3  'Birds  of  India,'  vol.  ii.  p.  96. 


DISPLAY  BY  THE  MALE.  397 

Sufficient  facts  have  now  been  given  to  show  with  what  care 
male  birds  display  their  various  charms,  and  this  they  do  with 
the  utmost  skill.  Whilst  preening  their  feathers,  they  have 
frequent  opportunities  for  admiring  themselves,  and  of  studying 
how  best  to  exhibit  their  beauty.  But  as  all  the  males  of  the 
same  species  display  themselves  in  exactly  the  same  manner,  it 
appears  that  actions,  at  first  perhaps  intentional,  have  become 
instinctive.  If  so,  we  ought  not  to  accuse  birds  of  conscious 
vanity;  yet  when  we  see  a  peacock  strutting  about,  with  ex- 
panded and  quivering  tail-feathers,  he  seems  the  very  emblem 
of  pride  and  vanity. 

The  various  ornaments  possessed  by  the  males  are  certainly 
of  the  highest  importance  to  them,  for  in  some  cases  they  have 
been  acquired  at  the  expense  of  greatly  impeded  powers  of  flight 
or  of  running.  The  African  night-jar  (Cosmetornis),  which  dur- 
ing the  pairing-season  has  one  of  its  primary  wing-feathers  de- 
veloped into  a  streamer  of  very  great  length,  is  thereby  much 
retarded  in  its  flight,  although  at  other  times  remarkable  for  its 
swiftness.  The  "unwieldy  size"  of  the  secondary  wing-feathers 
of  the  male  Argus  pheasant  are  said  "almost  entirely  to  deprive 
"the  bird  of  flight."  The  fine  plumes  of  male  birds  of  paradise 
trouble  them  during  a  high  wind.  The  extremely  long  tail- 
feathers  of  the  male  widow-birds  (Vidua)  of  Southern  Africa 
render  "their  flight  heavy;"  but  as  soon  as  these  are  cast  off 
they  fly  as  well  as  the  females.  As  birds  always  breed  when 
food  is  abundant,  the  males  probably  do  not  suffer  much  in- 
convenience in  searching  for  food  from  their  impeded  powers 
of  movement;  but  there  can  hardly  be  a  doubt  that  they  must 
be  much  more  liable  to  be  struck  down  by  birds  of  prey.  Nor 
can  we  doubt  that  the  long  train  of  the  peacock  and  the  long 
tail  and  wing-feathers  of  the  Argus  pheasant  must  render  them 
an  easier  prey  to  any  prowling  tiger-cat,  than  would  otherwise 
be  the  case.  Even  the  bright  colors  of  many  male  birds  cannot 
fail  to  make  them  conspicuous  to  their  enemies  of  all  kinds. 
Hence,  as  Mr.  Gould  has  remarked,  it  probably  is  that  such 
birds  are  generally  of  a  shy  disposition,  as  if  conscious  that  their 
beauty  was  a  source  of  danger,  and  are  much  more  difficult  to 
discover  or  approach,  than  the  somber  colored  and  compara- 
tively tame  females,  or  than  the  young  and  as  yet  unadorned 
males.''^ 


»3  On  the  Cosmetornis,  see  Livingstone's  'Expedition  to  the  Zam- 
besi,' I860,  p.  96.  On  the  Argus  pheasant,  Jardine's  'Nat.  Hist.  Lib.: 
Birds,'  vol.  xiv.  p.  167.  On  Birds  of  Paradise,  Lesson,  quoted  by  Brehm, 
'Thierleben,'  B.  iii.  s.  325,  On  the  widow-bird,  Barrow's  'Travels  in 
Africa,'  vol.  i.  p.  243,  and  'Ibis,'  vol.  iii.  1861,  p.  133.  Mr.  Gould,  on  the 
shyness  of  male  birds,  'Handbook  to  Birds  of  Australia,'  vol.  i.  1865, 
pp.  210,  457. 


398  THE   DESCENT  OF  MAN. 

It  is  a  more  curious  fact  that  the  males  of  some  birds  which 
are  provided  with  special  weapons  for  battle,  and  which  in  a 
state  of  nature  are  so  pugnacious  that  they  often  kill  each  other, 
suffer  from  possessing  certain  ornaments.  Cock-fighters  trim  the 
hackles  and  cut  off  the  combs  and  gills  of  their  cocks;  and  the 
birds  are  then  said  to  be  dubbed.  An  undubbed  bird,  as  Mr. 
Tegetmeier  insists,  "is  at  a  fearful  disadvantage;  the  comb  and 
"gills  offer  an  easy  hold  to  his  adversary's  beak,  and  as  a  cock 
"always  strikes  where  he  holds,  when  once  he  has  seized  his  foe, 
"he  has  him  entirely  in  his  power.  Even  supposing  that  the 
"bird  is  not  killed,  the  loss  of  blood  suffered  by  an  undubbed 
"cock  is  much  greater  than  that  sustained  by  one  that  has  been 
"trimmed. "°*  Young  turkey-cocks  in  fighting  always  seize  hold 
of  each  other's  wattles;  and  I  presume  that  the  old  birds  fight  in 
the  same  manner.  It  may  perhaps  be  objected  that  the  comb 
and  wattles  are  not  ornamental,  and  cannot  be  of  service  to  the 
birds  in  this  way;  but  even  to  our  eyes,  the  beauty  of  the  glossy 
black  Spanish  cock  is  much  enhanced  by  his  white  face  and 
crimson  comb;  and  no  one  who  has  ever  seen  the  splendid  blue 
wattles  of  the  male  Tragopan  pheasant,  distended  in  courtship, 
can  for  a  moment  doubt  that  beauty  is  the  object  gained.  From 
the  foregoing  facts  we  clearly  see  that  the  plumes  and  other 
ornaments  of  the  males  must  be  of  the  highest  importance  to 
them;  and  we  further  see  that  beauty  is  even  sometimes  more 
important  than  success  in  battle. 

9*  Tegetmeier,   'The  Poultry  Book,'  1866,  p.  139. 


LENGTH  OF  COURTSHIP.  399 


CHAPTER   XIV.. 

BIRDS— Continued. 

Choice  exerted  by  the  female— Length  of  courtship— "Unpaired  birds- 
Mental  qualities  and  taste  for  the  beautiful— Preference  or  antip- 
athy shown  by  the  female  for  particular  males— Variability  of  birds 
— ^Variations  sometimes  abrupt — Laws  of  variation— Formation  of 
ocelli— Gradations  of  character— Case  of  peacock,  Argus  pheasant, 
and  Urosticte. 

When  the  sexes  differ  in  beauty,  or  in  the  power  of  singing,  or 
in  producing  what  I  have  called  instrumental!  music,  it  is  almost 
invariably  the  male  who  surpasses  the  female.  These  qualities, 
as  we  have  just  seen,  are  evidently  of  high  importance  to  the 
male.  When  they  are  gained  for  only  a  part  of  the  year  it  is 
always  before  the  breeding  season.  It  is  the  male  alone  who 
elaborately  displays  his  varied  attractions,  and  often  performs 
strange  antics  on  the  ground  or  in  the  air,  in  the  presence  of  the 
female.  Each  male  drives  away,  or  if  he  can,  kills  his  rivals. 
Hence  we  may  conclude,  that  it  is  the  object  of  the  male  to  induce- 
the  female  to  pair  with  him,  and  for  this  purpose  he  tries  to  ex- 
cite or  charm  her  in  various  ways;  and  this  is  the  opinion  of  ail 
those  who  have  carefully  studied  the  habits  of  living  birds.  But 
there  remains  a  question  which  has  an  all-important  bearing  on 
sexual  selection,  namely,  does  every  male  of  the  same  species 
excite  and  attract  the  female  equally?  Or  does  she  exert  a  choice, 
and  prefer  certain  males?  This  latter  question  can  be  answered 
in  the  affirmative  by  much  direct  and  indirect  evidence.  It  is  far 
more  difficult  to  decide  what  qualities  determine  the  choice  of  the 
females;  but  here  again  we  have  some  direct  and  indirect  evi- 
dence that  it  is  to  a  large  extent  the  external  attractions  of  the 
male;  though  no  doubt  his  vigor,  courage,  and  other  mental 
qualities  come  into  play.  We  will  begin  with  the  indirect  evi- 
dence. 

Length  of  Courtship. — The  lengthened  period  during  which 
both  sexes  of  certain  birds  meet  day  after  day  at  an  appointed 
place  probably  depends  partly  on  the  courtship  being  a  prolonged 
affair,  and  partly  on  reiteration  in  the  act  of  pairing.    Thus  in 


400  THE  DESCENT  OF  MAJ^T. 

Germany  and  Scandinavia  the  balzing  or  leks  of  the  black-cocks 
last  from  the  middle  of  March,  all  through  April  into  May.  As 
many  as  forty  or  fifty,  or  even  more  birds  congregate  at  the  leks; 
and  the  same  place  is  often  frequented  during  successive  years. 
The  lek  of  the  capercailzie  lasts  from  the  end  of  March  to  the 
middle  or  even  end  of  May.  In  North  America  "the  partridge 
"dances"  of  the  Tetrao  phasianellus  "lasts  for  a  month  or  more." 
Other  kinds  of  grouse,  both  in  North  America  and  Eastern  Si- 
beria,^ follow  nearly  the  same  habits.  The  fowlers  discover  the 
hillocks  where  the  ruffs  congregate  by  the  grass  being  trampled 
bare,  and  this  shows  that  the  same  spot  is  long  frequented.  The 
Indians  of  Guiana  are  well  acquainted  with  the  cleared  arenas, 
where  they  expect  to  find  the  beautiful  cocks  of  the  Rock;  and 
the  natives  of  New  Guinea  know  the  trees  where  from  ten  to 
twenty  male  birds  of  paradise  in  full  plumage  congregate.  In  this 
latter  case  it  is  not  expressly  stated  that  the  females  meet  on  the 
same  trees,  but  the  hunters,  if  not  specially  asked,  would  probably 
not  mention  their  presence,  as  their  skins  are  valueless.  Small 
parties  of  an  African  weaver  (Ploceus)  congregate,  during  the 
breeding-season,  and  perform  for  hours  their  graceful  evolutions. 
Large  numbers  of  the  Solitary  snipe  (Scolopax  major)  assemble 
during  dusk  in  a  morass;  and  the  same  place  is  frequented  for 
the  same  purpose  during  successive  years;  here  they  may  be 
seen  running  about  "like  so  many  large  rats,"  puffing  out  their 
feathers,  flapping  their  wings,  and  uttering  the  strangest  cries.^ 
Some  of  the  above  birds, — ^the  black-cock,  capercailzie,  pheasant- 
grouse,  ruff.  Solitary  snipe,  and  perhaps  others, — are,  as  is  be- 
lieved, polygamists.  With  such  birds  it  might  have  been  thought 
that  the  stronger  males  would  simply  have  driven  away  the  weak- 
er, and  then  at  once  have  taken  possession  of  as  many  females 
as  possible;  but  if  it  be  indispensable  for  the  male  to  excite  or 
please  the  female,  we  can  understand  the  length  of  the  courtship 
and  the  congregation  of  so  many  individuals  of  both  sexes  at 
the  same  spot.  Certain  strictly  monogamous  species  likewise 
hold  nuptial  assemblages;  this  seems  to  be  the  case  in  Scandi- 
navia with  one  of  the  ptarmigans,  and  their  leks  last  from  the 

1  Nordman  describes  ('Bull.  Soc.  Imp.  des  Nat.  Moscou,'  1861,  torn. 
xxxiv.  p.  264)  the  balzen  of  Tetrao  urogalloides  in  Amur  Land.  He 
estimated  the  number  of  birds  assembled  at  above  a  hundred,  not 
counting  the  females,  which  lie  hid  in  the  surrounding  bushes.  The 
noises  uttered  differ  from  those  of  T.  urogallus. 

2  With  respect  to  the  assemblages  of  the  above  named  grouse  see 
Brehm,  'Thierleben.'  B.  iv.  s.  350;  also  L.  Lloyd,  'Game  Birds  of 
Sweden,'  1867,  pp.  19,  78.  Richardson,  'Fauna  Bor.  Americana,'  Birds, 
p.  362.  References  in  regard  to  the  assemblages  of  other  birds  have 
already  been  given.  On  Paradisea  see  "Wallace,  in  'Annals  and  Mag. 
of  Nat.  Hist.'  vol.  xx.  1857,  p.  412.    On  the  snipe,  Lloyd,  ibid.  p.  221. 


UNPAIRED  BIRDS.  401 

middle  of  March  to  the  middle  of  May.  In  Australia  the  lyre- 
bird (Menura  superba)  forms  "small  round  hillocks,"  and  the  M. 
Albert!  scratches  for  itself  shallow  holes,  or,  as  they  are  called 
by  the  natives,  corroborying  places,  where  it  is  believed  both 
sexes  assemble.  The  meetings  of  the  M.  superba  are  sometimes 
very  large;  and  an  account  has  lately  been  published^  by  a 
traveler,  who  heard  in  a  valley  beneath  him,  thickly  covered 
with  scrub,  "a  din  which  completely  astonished"  him;  on  crawl- 
ing onwards  he  beheld  to  his  amazement  about  one  hundred  and 
fifty  of  the  magnificent  lyre-cocks,  "ranged  in  order  of  battle,  and 
"fighting  with  indescribable  fury."  The  bowers  of  the  Bower- 
birds  are  the  resort  of  both  sexes  during  the  breeding-season; 
and  "here  the  males  meet  and  contend  with  each  other  for  the 
"favors  of  the  female,  and  here  the  latter  assemble  and  coquet 
"with  the  males."  With  two  of  the  genera,  the  same  bower  is 
resorted  to  during  many  years.* 

The  common  magpie  (Corvus  pica,  Linn.),  as  I  have  been  in- 
formed by  the  Rev.  W.  Darwin  Fox,  used  to  assemble  from  all 
parts  of  Delamere  Forest,  in  order  to  celebrate  the  "great  mag- 
"pie  marriage."  Some  years  ago  these  birds  abounded  in  extraor- 
dinary numbers,  so  that  a  gamekeeper  killed  in  one  morning  nine- 
teen males,  and  another  killed  by  a  single  shot  seven  birds  at 
roost  together.  They  then  had  the  habit  of  assembling  very 
early  in  the  spring  at  particular  spots,  where  they  could  be  seen 
in  flocks,  chattering,  sometimes  fighting,  bustling  and  fiying  about 
the  trees.  The  whole  affair  was  evidently  considered  by  the  birds 
as  one  of  the  highest  importance.  Shortly  after  the  meeting  they 
all  separated,  and  were  then  observed  by  Mr.  Fox  and  others  to 
be  paired  for  the  season.  In  any  district  in  which  a  species  does 
not  exist  in  large  numbers,  great  assemblages  cannot,  of  course, 
be  held,  and  the  same  species  may  have  different  habits  in  dif- 
ferent countries.  For  instance,  I  have  heard  of  only  one  instance, 
from  Mr.  Wedderburn,  of  a  regular  assemblage  of  black  game  in 
Scotland,  yet  these  assemblages  are  so  well  known  in  Germany 
and  Scandinavia  that  they  have  received  special  names. 

Unpaired  Birds. — From  the  facts  now  given,  we  may  conclude 
that  the  courtship  of  birds,  belonging  to  widely  different  groups, 
is  often  a  prolonged,  delicate,  and  troublesome  affair.  There  is 
even  reason  to  suspect,  improbable  as  this  will  at  first  appear, 
that  some  males  and  females  of  the  same  species,  inhabiting  the 
same  district,  do  not  always  please  each  other,  and  consequently 
do  not  pair.     Many  accounts  have  been  published  of  either  the 

8  Quoted  by  Mr.  T.  W.  Wood  in  the  'Student,'  April,  1870,  p.  125. 
*  Gould,  'Handbook  to  the  Birds  of  Australia,'  vol.  i.  pp.  300,  308,  448, 
451.    On  the  ptarmigan,  above  alluded  to,  see  Lloyd,  ibid.  p.  129. 
27 


402  THE  DESCENT  OF  MAN. 

male  or  female  of  a  pair  having  been  shot,  and  quickly  replaced 
by  another.  This  has  been  observed  more  frequently  with  the 
magpie  than  with  any  other  bird,  owing  perhaps  to  its  conspicu- 
ous appearance  and  nest.  The  illustrious  Jenner  states  that  in 
Wiltshire  one  of  a  pair  was  daily  shot  no  less  than  seven  times 
successively,  "but  all  to  no  purpose,  for  the  remaining  magpie 
**soon  found  another  mate;"  and  the  last  pair  reared  their  young. 
A  new  partner  is  generally  found  on  the  succeeding  day;  but 
Mr.  Thompson  gives  the  case  of  one  being  replaced  on  the  even- 
ing of  the  same  day.  Even  after  the  eggs  are  hatched,  if  one  of 
the  old  birds  is  destroyed  a  mate  will  often  be  found;  this  oc- 
curred after  an  interval  of  two  daj^s,  in  a  case  recently  observed 
by  one  of  Sir  J.  Lubbock's  keepers.*  The  first  and  most  obvious 
conjecture  is  that  male  magpies  must  be  much  more  numerous 
than  females;  and  that  in  the  above  cases,  as  well  as  in  many 
others  which  could  be  given,  the  males  alone  had  been  killed. 
This  apparently  holds  good  in  some  instances,  for  the  game- 
keepers in  Delamere  Forest  assured  Mr.  Fox  that  the  magpies 
and  carrion-crows  which  they  formerly  killed  in  succession  in 
large  numbers  near  their  nests,  were  all  males;  and  they  ac- 
counted for  this  fact  by  the  males  being  easily  killed  whilst  bring- 
ing food  to  the  sitting  females.  Macgillivray,  however,  gives, 
on  the  authority  of  an  excellent  observer,  an  instance  of  three 
magpies  successively  killed  on  the  same  nest,  which  were  all  fe- 
males; and  another  case  of  six  magpies  successively  killed  whilst 
sitting  on  the  same  eggs,  which  renders  it  probable  that  most  of 
them  were  females;  though,  as  I  hear  from  Mr.  Fox,  the  male 
will  sit  on  the  eggs  when  the  female  is  killed. 

Sir  J.  Lubbock's  gamekeeper  has  repeatedly  shot,  but  how 
often  he  could  not  say,  one  of  a  pair  of  jays  (Garrulus  glandarius), 
and  has  never  failed  shortly  afterwards  to  find  the  survivor  re- 
matched.  Mr.  Fox,  Mr.  F.  Bond,  and  others  have  shot  one  of 
a  pair  of  carrion-crows  (Corvus  corone),  but  the  nest  was  soon 
again  tenanted  by  a  pair.  These  birds  are  rather  common;  but 
the  peregrine-falcon  (Falco  peregrinus)  is  rare,  yet  Mr.  Thompson 
states  that  in  Ireland  "if  either  an  old  male  or  female  be  killed 
"in  the  breeding  season  (not  an  uncommon  circumstance),  an- 
"other  mate  is  found  within  a  very  few  days,  so  that  the  eyries, 
"notwithstanding  such  casualties,  are  sure  to  turn  out  their  com- 
"plement  of  young."  Mr.  Jenner  Weir  has  known  the  same  thing 
with  the  peregrine-falcons  at  Beachy  Head.  The  same  observer 
informs  me  that  three  kestrels  (Falco  tinnunculus),  all  males, 
were  killed  one  after  the  other  whilst  attending  the  same  nest; 

^  On  magpies,  Jenner,  in  'Phil.  Transact.'  1824,  p.  21.  Macg-illivray, 
'Hist.  British  Birds,'  vol.  i.  p.  570.  Thompson,  in  'Annals  and  Mag-,  of 
Nat.  Hist.'  vol.  viii.  1842,  p.  494. 


UNPAIRED  BIRDS.  403 

two  of  these  were  in  mature  plumage,  but  the  third  was  in  the 
plumage  of  the  previous  year.  Even  with  the  rare  golden  eagle 
(Aquila  chrysaetos),  Mr.  Birkbeck  was  assured  by  a  trustworthy 
gamekeeper  in  Scotland,  that  if  one  is  killed,  another  is  soon 
found.  So  with  the  white  owl  (Strix  flammea),  "the  survivor 
"readily  found  a  mate,  and  the  mischief  went  on." 

White  of  Selborne,  who  gives  the  case  of  the  owl,  adds  that 
he  knew  a  man,  who  from  believing  that  patridges  when  paired 
were  disturbed  by  the  males  fighting,  used  to  shoot  them;  and 
though  he  had  widowed  the  same  female  several  times,  she  always 
soon  found  a  fresh  partner.  This  same  naturalist  ordered  the 
sparrows,  which  deprived  the  house-martins  of  their  nests,  to 
be  shot;  but  the  one  which  was  left,  "be  it  cock  or  hen,  presently 
"procured  a  mate,  and  so  for  several  times  following."  I  could 
add  analogous  cases  relating  to  the  chaffinch,  nightingale,  and 
redstart.  With  respect  to  the  latter  bird  (Phsenicura  ruticilla),  a 
writer  expresses  much  surprise  how  the  sitting  female  could  so 
soon  have  given  effectual  notice  that  she  was  a  widow,  for  the 
species  was  not  common  in  the  neighborhood.  Mr.  Jenner  Weir 
has  mentioned  to  me  a  nearly  similar  case;  at  Blackheath  he 
never  sees  or  hears  the  note  of  the  wild  bullfinch,  yet  when  one 
of  his  caged  males  has  died,  a  wild  one  in  the  course  of  a  few 
days  has  generally  come  and  perched  near  the  wi'dowed  female, 
whose  call-note  is  not  loud.  I  will  give  only  one  other  fact,  on 
the  authority  of  this  same  observer;  one  of  a  pair  of  starlings 
(Sturnus  vulgaris)  was  shot  in  the  morning;  by  noon  a  new 
mate  was  found;  this  was  again  shot,  but  before  night  the  pair 
was  complete;  so  that  the  disconsolate  widow  or  widower  was 
thrice  consoled  during  the  same  day.  Mr.  Engleheart  also  in- 
forms me  that  he  used  during  several  years  to  shoot  one  of  a  pair 
of  starlings  which  built  in  a  hole  in  a  house  at  Blackheath;  but 
the  loss  was  always  immediately  repaired.  During  one  season 
he  kept  an  account,  .and  found  that  he  had  shot  thirty-five  birds 
from  the  same  nest;  these  consisted  of  both  males  and  fe- 
males, but  in  what  proportion  he  could  not  say:  nevertheless, 
after  all  this  destruction,  a  brood  was  reared.^ 

These  facts  well  deserve  attention.  How  is  it  that  there  are 
birds  enough,  ready  to  replace  immediately  a  lost  mate  of  either 
sex?  Magpies,  jays,  carrion-crows,  partridges,  and  some  other 
birds,  are  always  seen  during  the  spring  in  pairs,  and  never  by 


«  On  the  peregrine  falcon  see  Thompson,  'Nat.  Hist,  of  Ireland, 
Birds,'  vol.  i,  1849,  p.  39.  On  owls,  sparrows,  and  partridges,  see  White, 
'Nat.  Hist,  of  Selborne,'  edit,  of  1825,  vol.  i.  p.  139.  On  the  Phoenicura, 
see  Loudon's  'Mag.  of  Nat.  Hist.'  vol.  vii.  1834,  p.  245.  Brehm  ('Thier- 
leben,'  B.  iv.  s.  991)  also  alludes  to  cases,  of  birds  thrice  mated  during 
the  same  day. 


404  THE  DESCENT  OP  MAN. 

themselves;  and  these  offer  at  first  sight  the  most  perplexing 
cases.  But  hirds  of  the  same  sex,  although  of  course  not  truly 
paired,  sometimes  live  in  pairs  or  in  small  parties,  as  is  known  to 
be  the  case  with  pigeons  and  partridges.  Birds  also  sometimes 
live  in  triplets,  as  has  been  observed  with  starlings,  carrion-crows, 
parrots  and  partridges.  With  partridges  two  females  have  been 
known  to  live  with  one  male,  and  two  males  with  one  female.  In 
all  such  cases  it  is  probable  that  the  union  would  be  easily  broken; 
and  one  of  the  three  would  readily  pair  with  a  widow  or  widower. 
The  males  of  certain  birds  may  occasionally  be  heard  pouring  forth 
their  love-song  long  after  the  proper  time,  showing  that  they 
have  either  lost  or  never  gained  a  mate.  Death  from  accident 
or  disease  of  one  of  a  pair,  would  leave  the  other  free  and  single; 
and  there  is  reason  to  believe  that  female  birds  during  the 
breeding-season  are  especially  liable  to  premature  death.  Again, 
birds  which  have  had  their  nests  destroyed,  or  barren  pairs,  or  re- 
tarded individuals,  would  easily  be  induced  to  desert  their  mates, 
and  would  probably  be  glad  to  take  what  share  they  could  of  the 
pleasures  and  duties  of  rearing  offspring  although  not  their  own.' 
Such  contingencies  as  these  probably  explain  most  of  the  fore- 
going cases.®  Nevertheless,  it  is  a  strange  fact  that  within  the 
same  district,  during  the  height  of  the  breeding-season,  there 
should  be  so  many  males  and  females  always  ready  to  repair  the 
loss  of  a  mated  bird.  Why  do  not  such  spare  birds  immediately 
pair  together?  Have  we  not  some  reason  to  suspect,  and  the  sus- 
picion has  occurred  to  Mr.  Jenner  Weir,  that  as  the  courtship  of 


■y  See  White  ('Nat.  Hist,  of  Selborne,'  1825,  vol.  i.  p.  140)  on  the  exist- 
ence, early  in  the  season,  of  small  coveys  of  male  partridges,  of  which 
fact  I  have  heard  other  instances.  See  Jenner,  on  the  retarded  state 
of  the  generative  organs  in  certain  birds,  in  'Phil.  Transact.'  1824.  In 
regard  to  birds  living  in  triplets,  I  owe  to  Mr.  Jenner  Weir  the  cases 
of  the  starlings  and  parrots,  and  to  Mr.  Fox,  of  partridges:  on  car- 
rion-crows, see  the  'Field,'  1868,  p.  415.  On  various  male  birds  singing 
after  the  proper  period,  see  Rev.  L..  Jenyns,  'Observations  in  Natural 
History,'  1846.  p.  87. 

8  The  following  case  has  been  given  ('The  Times,'  Aug.  6th,  1868)  by 
the  Rev.  F.  O.  Morris,  on  the  authority  of  the  Hon.  and  Rev.  O.  W. 
Forester.  "The  gamekeeper  here  found  a  hawk's  nest  this  year,  with 
"five  young  ones  on  it.  He  took  four  and  killed  them,  but  left  one 
"with  its  wings  clipped  as  a  decoy  to  destroy  the  old  ones  by.  They 
"were  both  shot  next  day,  in  the  act  of  feeding  the  young  one,  and 
"the  keeper  thought  it  was  done  with.  The  next  day  he  came  again 
"and  found  two  other  charitable  hawks,  who  had  come  with  an  adopted 
"feeling  to  succor  the  orphan.  These  two  he  killed,  and  then  left  the 
"nest.  On  returning  afterwards  he  found  two  more  charitable  in- 
"dlviduals  on  the  some  errand  of  mercy.  One  of  these  he  killed;  the 
"other  he  also  shot,  but  could  not  find.  No  more  came  on  the  like 
"fruitless  errand." 


MENTAL  QUALITIES.  405 

birds  appears  to  be  in  many  cases  prolonged  and  tedious,  so  it 
occasionally  happens  that  certain  males  and  females  do  not  suc- 
ceed during  the  proper  season,  in  exciting  each  other's  love,  and 
consequently  do  not  pair?  This  suspicion  will  appear  somewhat 
less  improbable  after  we  have  seen  what  strong  antipathies  and 
preferences  female  birds  occasionally  evince  towards  particular 
males. 

Mental  Qualities  of  Birds,  and  their  Taste  for  the  Beautiftil. — 
Before  we  further  discuss  the  question  whether  the  females  select 
the  more  attractive  males  or  accept  the  first  whom  they  may  en- 
counter, it  will  be  advisable  briefly  to  consider  the  mental  powers 
of  birds.  Their  reason  is  generally,  and  perhaps  justly,  ranked  as 
low;  yet  some  facts  could  be  given^  leading  to  an  opposite  conclu- 
sion. Low  powers  of  reasoning,  however,  are  compatible,  as  we 
see  with  mankind,  with  strong  affections,  acute  perception,  and  a 
taste  for  the  beautiful;  and  it  is  with  these  latter  qualities  that 
we  are  here  concerned.  It  has  often  been  said  that  parrots  be- 
come so  deeply  attached  to  each  other  that  when  one  dies  the 
other  pines  for  a  long  time;  but  Mr.  Jenner  Weir  thinks  that  with 
most  birds  the  strength  of  their  affection  has  been  much  exag- 
gerated. Nevertheless  when  one  of  a  pair  in  a  state  of  nature  has 
been  shot,  the  survivor  has  been  heard  for  days  afterwards  utter- 
ing a  plaintive  call;  and  Mr.  St.  John  gives  various  facts  proving 
the  attachment  of  mated  birds.^*^  Mr.  Bennett  relates^^  that  in 
China  after  a  drake  of  the  beautiful  mandarin  Teal  had  been 
stolen,  the  duck  remained  disconsolate,  though  sedulously  courted 
by  another  mandarin  drake,  who  displayed  before  her  all  his 
charms.  After  an  interval  of  three  weeks  the  stolen  drake  was 
recovered,  and  instantly  the  pair  recognized  each  other  with  ex- 
treme joy.  On  the  other  hand  starlings,  as  we  have  seen,  may  be 
consoled  thrice  in  the  same  day  for  the  loss  of  their  mates.  Pig- 
eons have  such  excellent  local  memories,  that  they  have  been 
known  to  return  to  their  former  homes  after  an  interval  of  nine 

»  I  am  indebted  to  Prof.  Newton  for  the  following-  passage  from  Mr. 
Adam's  'Travels  of  a  Naturalist,'  1870,  p.  278.  Speaking  of  Japanese 
nut-hatches  in  confinement  he  says".  "Instead  of  the  more  yielding 
"fruit  of  the  yew,  which  is  the  usual  food  of  the  nut-hatch  of  Japan, 
"at  one  time  I  substituted  hard  hazel-nuts.  As  the  bird  was  unable 
"to  crack  them,  he  placed  them  one  by  one  in  his  water-glass,  evi- 
"dently  with  the  notion  that  they  would  in  time  become  softer— an 
"interesting  proof  of  intelligence  on  the  part  of  these  birds." 

10  «A  Tour  in  Sutherlandshire,'  vol.  i.  1849,  p.  185.  Dr.  Buller  says 
('Birds  of  New  Zealand,'  1872,  p.  56)  that  a  male  King  Lory  was  killed: 
and  the  female  "fretted  and  moped,  refused  her  food,  and  died  of  a 
"broken  heart." 

^  'Wanderings  in  New  South  Wales,'  vol.  ii.  18:^4,  p.  62. 

27 


406  THE  DESCENT  OP  MAN. 

months,  yet,  as  I  hear  from  Mr.  Harrison  Weir,  if  a  pair  which 
naturally  would  remain  mated  for  life  be  separated  for  a  few 
weeks  during  the  winter,  and  afterwards  matched  with  other 
birds,  the  two,  when  brought  together  again,  rarely,  if  ever,  rec- 
ognize each  other. 

Birds  sometimes  exhibit  benevolent  feelings;  they  will  feed  the 
deserted  young  ones  even  of  distinct  species,  but  this  perhaps 
ought  to  be  considered  as  a  mistaken  instinct.  They  will  feed, 
as  shown  in  an  earlier  part  of  this  work,  adult  birds  of  their  own 
species  which  have  become  blind.  Mr.  Buxton  gives  a  curious 
account  of  a  parrot  which  took  care  of  a  frost-bitten  and  crippled 
bird  of  a  distinct  species,  cleansed  her  feathers,  and  defended  her 
from  the  attacks  of  the  other  parrots  which  roamed  freely  about 
his  garden.  It  is  a  still  more  curious  fact  that  these  birds  ap- 
parently evince  some  sympathy  for  the  pleasures  of  their  fellows. 
"When  a  pair  of  cockatoos  made  a  nest  in  an  acacia  tree,  "it  was 
"ridiculous  to  see  the  extravagant  interest  taken  in  the  matter  by 
"the  others  of  the  same  species."  These  parrots,  also  evinced  un- 
bounded curiosity,  and  clearly  had  "the  idea  of  property  and  pos- 
"session."^^  They  have  good  memories,  for  in  the  Zoological 
Gardens  they  have  plainly  recognized  their  former  masters  after 
an  interval  of  some  months. 

Birds  possess  acute  powers  of  observation.  Every  mated  bird, 
of  course,  recognizes  its  fellow.  Audubon  states  that  a  certain 
number  of  mocking-thrushes  (Mimus  polyglottus)  remain  all  the 
year  round  in  Louisiana,  whilst  others  migrate  to  the  Eastern 
States;  these  latter,  on  their  return,  are  instantly  recognized,  and 
always  attacked,  by  their  southern  brethren.  Birds  under  con- 
finement distinguish  different  persons,  as  is  proved  by  the  strong 
and  permanent  antipathy  or  affection  which  they  show,  without 
any  apparent  cause,  towards  certain  individuals.  I  have  heard 
of  numerous  instances  with  jays,  partridges,  canaries,  and  es- 
pecially bullfinches.  Mr.  Hussey  has  described  in  how  extraor- 
dinary a  manner  a  tamed  partridge  recognized  everybody;  and  its 
likes  and  dislikes  vvrere  very  strong.  This  bird  seemed  "fond  of 
"gay  colors,  and  no  new  gown  or  cap  could  be  put  on  without 
"catching  his  attention."^^  Mr.  Hewitt  has  described  the  habits 
of  some  ducks  (recently  descended  from  wild  birds),  which,  at  the 
approach  of  a  strange  dog  or  cat,  would  rush  headlong  into  the 
water,  and  exhaust  themselves  in  their  attempts  to  escape;  but 
they  knew  Mr.  Hewitt's  own  dogs  and  cats  so  well,  that  they 
would  lie  down  and  bask  in  the  sun  close  to  them.  They  always 
moved  away  from  a  strange  man,  and  so  they  would  from  the 


^  'Acclimatization   of   Parrots,'   by  C.   Buxton,    M.    P.    'Annals   and 
Mag.  of  Nat.  Hist.'  Nov.  1868,  p.  381. 
13  'The  ZoolOt?ist,'  1847-1848,  p.  1602. 


*  MENTAL   QUALITIES.  407 

lady  who  attended  them,  if  she  made  any  great  change  in  her 
dress.  Audubon  relates  that  he  reared  and  tamed  a  wild  turkey 
which  always  ran  away  from  any  strange  dog;  this  bird  escaped 
into  the  woods,  and  some  days  afterwards  Audubon  saw,  as  he 
thought,  a  wild  turkey,  and  made  his  dog  chase  it;  but  to  his 
astonishment,  the  bird  did  not  run  away,  and  the  dog,  when  he 
came  up,  did  not  attack  the  bird,  for  they  mutually  recognized 
each  other  as  old  friends.^* 

Mr.  Jenner  Weir  is  convinced  that  birds  pay  particular  atten- 
tion to  the  colors  of  other  birds,  sometimes  out  of  jealousy,  and 
sometimes  as  a  sign  of  kinship.  Thus  he  turned  a  reed-bunting 
(Emberiza  schceniculus),  which  had  acquired  its  black  head-dress, 
into  his  aviary,  and  the  new-comer  was  not  noticed  by  any  bird, 
except  by  a  bullfinch,  which  is  likewise  black-headed.  This  bull- 
finch was  a  very  quiet  bird,  and  had  never  before  quarreled  with 
any  of  its  comrades,  including  another  reed-bunting,  which  had 
not  as  yet  become  black-headed:  but  the  reed-bunting  with  a 
black  head  was  so  unmercifully  treated,  that  it  had  to  be  re- 
moved. Spiza  cyanea,  during  the  breeding-season,  is  of  a  bright 
blue  color;  and  though  generally  peaceable,  it  attacked  S.  ciris, 
which  has  only  the  head  blue,  and  completely  scalped  the  unfortu- 
nate bird.  Mr.  Weir  was  also  obliged  to  turn  out  a  robin,  as  it 
fiercely  attacked  all  the  birds  in  his  aviary  with  any  red  in  their 
plumage,  but  no  other  kinds;  it  actually  killed  a  red-breasted 
crossbill,  and  nearly  killed  a  goldfinch.  On  the  other  hand,  he  has 
observed  that  some  birds,  when  first  introduced,  fly  towards  the 
species  which  resemble  them  most  in  color,  and  settle  by  their 
sides. 

As  male  birds  display  their  fine  plumage  and  other  ornaments 
with  so  much  care  before  the  females,  it  is  obviously  probable  that 
these  appreciate  the  beauty  of  their  suitors.  It  is,  however,  dif- 
ficult to  obtain  direct  evidence  of  their  capacity  to  appreciate 
beauty.  When  birds  gaze  at  themselves  in  a  looking-glass  (of 
which  many  instances  have  been  recorded)  we  cannot  feel  sure 
that  it  is  not  from  jealousy  of  a  supposed  rival,  though  this  is  not 
the  conclusion  of  some  observers.  In  other  cases  it  is  diflicult  to 
distinguish  between  mere  curiosity  and  admiration.  It  is  per- 
haps the  former  feeling  which,  as  stated  by  Lord  Lilford,^^  at- 
tracts the  ruff  towards  any  bright  object,  so  that,  in  the  Ionian  Is- 
lands, "it  will  dart  down  to  a  bright-colored  handkerchief,  regard- 
"less  of  repeated  shots."  The  common  lark  is  drawn  down  from 
the  sky,  and  is  caught  in  large  numbers,  by  a  small  mirror  made 


"  Hewitt  on  wild  ducks,  'Journal  of  Horticulture,'  Jan.  13,  1863,  p.  39. 
Audubon  on  the  wild  turkey,  'Ornith.  Biography,'  vol.  i.  p.  14.  On 
the  mocking-thrush,  ibid.  vol.  i.  p.  110. 

IB  The  'Ibis,'  vol.  ii.  1860,  p.  344, 


408  THE   DESCENT  OF  MAN. 

to  move  and  glitter  In  the  sun.  Is  it  admiration  or  curiosity  which 
leads  the  magpie,  raven,  and  some  other  birds  to  steal  and  secrete 
bright  objects,  such  as  silver  articles  or  jewels? 

Mr.  Gould  states  that  certain  hrmming-birds  decorate  the  out- 
sides  of  their  nests  "with  the  utmost  taste;  they  instinctively  fas- 
"ten  thereon  beautiful  pieces  of  flat  lichen,  the  larger  pieces  in  the 
"middle,  and  the  smaller  on  the  part  attached  to  the  branch. 
"Now  and  then  a  pretty  feather  is  intertwined  or  fastened  to  the 
"outer  sides,  the  stem  being  always  so  placed  that  the  feather 
"stands  out  beyond  the  surface."  The  best  evidence,  however,  of  a 
taste  for  the  beautiful  is  afforded  by  the  three  genera  of  Australian 
bower-birds  already  mentioned.  Their  bowers  (see  fig.  46,  p.  378), 
where  the  sexes  congregate  and  play  strange  antics,  are  variously 
constructed,  but  what  most  concerns  us  is,  that  they  are  decorated 
by  the  several  species  in  a  different  manner.  The  Satin  bower- 
bird  collects  gaily-colored  articles,  such  as  the  blue  tail-feathers  of 
parrakeets,  bleached  bones  and  shells,  which  it  sticks  between  the 
twigs,  or  arranges  at  the  entrance.  Mr.  Gould  found  in  one  bower 
a  neatly-worked  stone  tomahawk  and  a  slip  of  blue  cotton,  evi- 
dently procured  from  a  native  encampment.  These  objects  are 
continually  re-arranged,  and  carried  about  by  the  birds  whilst  at 
play.  The  bower  of  the  Spotted  bower-bird  "is  beautifully  lined 
"with  tall  grasses,  so  disposed  that  the  heads  nearly  meet,  and 
"the  decorations  are  very  profuse."  Round  stones  are  used  to 
keep  the  grass-stems  in  their  proper  places,  and  to  make  diver- 
gent paths  leading  to  the  bower.  The  stones  and  shells  are  often 
brought  from  a  great  distance.  The  Regent  bird,  as  described  by 
Mr.  Ramsay,  ornaments  its  short  bower  with  bleached  land-shells 
belonging  to  five  or  six  species,  and  with  "berries  of  various  col- 
"ors,  blue,  red,  and  black,  which  give  it  when  tresh,  a  very  pretty 
"appearance.  Besides  these  there  were  several  newly-picked  leaves 
"and  young  shoots  of  a  pinkish  color,  the  whole  showing  a  decided 
"taste  for  the  beautiful."  Well  may  Mr.  Gould  say,  that  these 
"highly  decorated  halls  of  assembly  must  be  regarded  as  the  most 
"wonderful  instances  of  bird-architecture  yet  discovered;"  and  the 
taste,  as  we  see,  of  the  several  species  certainly  differs.^^ 

Preference  for  particular  Males  hy  the  Females.  — Having  made 
these  preliminary  remarks  on  the  discrimination  and  taste  of 
birds,  I  will  give  all  the  facts  known  to  me,  which  bear  on  the  pref- 
erence shown  by  the  female  for  particular  males.  It  is  certain 
that  distinct  species  of  birds  occasionally  pair  in  a  state  of  nature 

^®  On  the  ornamented  nests  of  humming'-birds,  Gould,  'Introduction 
to  the  Trochilidae,'  1861,  p.  19.  On  the  bower-birds,  Gould,  'Handbook 
to  the  Birds  of  Australia,'  1865,  vol.  i.  pp.  444-461.  Ramsay,  in  the  'Ibis,' 
1867,  p.  456. 


PREFERENCE  BY  THE  FEMALE.  409 

and  produce  hybrids.  Many  instances  could  be  given:  thus  Mac- 
gillivray  relates  how  a  male  blackbird  and  female  thrush  "fell  in 
"love  with  each  other,"  and  produced  offspring.^^  Several  years 
ago  eighteen  cases  had  been  recorded  of  the  occurrence  in  Great 
Britain  of  hybrids  between  the  black  grouse  and  pheasant  ;^^  but 
most  of  these  cases  may  perhaps  be  accounted  for  by  solitary  birds 
not  finding  one  of  their  own  species  to  pair  with.  With  other 
birds,  as  Mr.  Jenner  Weir  has  reason  to  believe,  hybrids  are  some- 
times the  result  of  the  casual  intercourse  of  birds  building  in  close 
proximity.  But  these  remarks  do  not  apply  to  the  many  recorded 
instances  of  tamed  or  domestic  birds,  belonging  to  distinct  species, 
which  have  become  absolutely  fascinated  with  each  other,  although 
living  with  their  own  species.  Thus  Waterton^^  states  that  out  of 
a  flock  of  twenty-three  Canada  geese,  a  female  paired  with  a  soli- 
tary Bernicle  gander,  although  so  different  in  appearance  and  size; 
and  they  produced  hybrid  offspring,  A  male  widgeon  (Mareca 
penelope),  living  with  females  of  the  same  species,  has  been 
known  to  pair  with  a  pintail  duck,  Querciuedula  acuta.  Lloyd  de- 
scribes the  remarkable  attachment  between  a  shield-drake  (Tad- 
orna  vulpanser)  and  a  common  duck.  Many  additional  instances 
could  be  given;  and  the  Rev.  B.  S.  Dixon  remarks  that  "those 
"who  have  kept  many  different  species  of  geese  together,  well 
"know  what  unaccountable  attachments  they  are  frequently  form- 
"ing,  and  that  they  are  quite  as  likely  to  pair  and  rear  young  with 
"individuals  of  a  race  (species)  apparently  the  most  alien  to 
"themselves,  as  with  their  own  stock." 

The  Rev.  W.  D.  Fox  informs  me  that  he  possessed  at  the  same 
time  a  pair  of  Chinese  geese  (Anser  cygnoides),  and  a  common 
gander  with  three  geese.  The  two  lots  kept  quite  separate,  until 
the  Chinese  gander  seduced  one  of  the  common  geese  to  live  with 
him.  Moreover,  of  the  young  birds  hatched  from  the  eggs  of  the 
common  geese,  only  four  were  pure,  the  other  eighteen  proving 
hybrids;  so  that  the  Chinese  gander  seems  to  have  had  prepotent 
charms  over  the  common  gander.  I  will  give  only  one  other  case; 
Mr.  Hewitt  states  that  a  wild  duck,  reared  in  captivity,  "after 
"breeding  a  couple  of  seasons  with  her  own  mallard,  at  once  shook 
"him  off  on  my  placing  a  male  Pintail  on  the  water.  It  was  evi- 
"dently  a  case  of  love  at  first  sight,  for  she  swam  about  the  new- 

"  'Hist,  of  British  Birds,'  vol.  ii.  p.  92. 

18  'Zoolog-ist,'  1853-1S54,  p.  8946. 

i»  Waterton,  'Essays  on  Nat.  Hist.'  2nd  series,  pp.  42  and  117.  For  the 
following  statements  see  on  the  widgeon,  Loudon's  'Mag-,  of  Nat.  Hist.' 
vol.  ix.  p.  616;  L.  Lloyd,  'Scandinavian  Adventures,'  vol.  i.  1854,  p.  452.- 
Dixon,  'Ornamental  and  Domestic  Poultry,'  p.  137;  Hewitt,  in  'Journal 
of  Horticulture,'  Jan.  13,  1863,  p.  40;  Bechstein,  'Stubenvogel,'  1840,  s.  2S0. 
Mr.  J.  Jenner  Weir  has  lately  given  me  an  analogous  case  with  ducks 
of  two  species. 


410  THE  DESCENT  OF  MAN. 

"comer  caressingly,  though  he  appeared  evidently  alarmed  and 
"averse  to  her  overtures  of  affection.  From  that  hour  she  forgot 
"her  old  partner.  Winter  passed  by,  and  the  next  spring  the  Pin- 
"tail  seemed  to  have  become  a  convert  to  her  blandishments,  tor 
"they  nested  and  produced  seven  or  eight  young  ones." 

What  the  charm  may  have  been  in  these  several  cases,  beyond 
mere  novelty,  we  cannot  even  conjecture.  Color,  however,  some- 
times comes  into  play;  for  in  order  to  raise  hybrids  from  the  sis- 
kin (Fringilla  spinus)  and  the  canary,  it  is  much  the  best  plan, 
according  to  Bechstein,  to  place  birds  of  the  same  tint  together, 
Mr.  Jenner  Weir  turned  a  female  canary  into  his  aviary, 
where  there  were  male  linnets,  goldfinches,  siskins,  greenfinches, 
chafiinches,  and  other  birds,  in  order  to  see  which  she  would 
choose;  but  there  never  was  any  doubt,  and  the  greenfinch  carried 
the  day.     They  paired  and  produced  hybrid  offspring. 

The  fact  of  the  female  preferring  to  pair  with  one  male  rather 
than  with  another  of  the  same  species,  is  not  so  likely  to  excite  at- 
tention, as  when  this  occurs,  as  we  have  just  seen,  between  dis- 
tinct species.  The  former  cases  can  best  be  observed  with  domes- 
ticated or  confined  birds;  but  these  are  often  pampered  by  high 
feeding,  and  sometimes  have  their  instincts  vitiated  to  an  extreme 
degree.  Of  this  latter  fact  I  could  give  sufficient  proofs  with  pig- 
eons, and  especially  with  fowls,  but  they  cannot  be  here  related. 
Vitiated  instincts  may  also  account  for  some  of  the  hybrid  unions 
above  mentioned;  but  in  many  of  these  cases  the  birds  v/ere  al- 
lowed to  range  freely  over  large  ponds,  and  there  is  no  reason  to 
suppose  that  they  were  unnaturally  stimulated  by  high  feeding. 

With  respect  to  birds  in  a  state  of  nature,  the  first  and  most 
obvious  supposition  which  will  occur  to  every  one  is  that  the 
female  at  the  proper  season  accepts  the  first  male  whom  she  may 
encounter;  but  she  has  at  least  the  opportunity  for  exerting  a 
choice,  as  she  is  almost  invariably  pursued  by  many  males. 
Audubon — and  we  must  remember  that  he  spent  a  long  life  in 
prowling  about  the  forests  of  the  United  States  and  observing 
the  birds — does  not  doubt  that  the  female  deliberately  chooses 
her  mate;  thus,  speaking  of  a  woodpecker,  he  says  the  hen  is  fol- 
lowed by  half-a-dozen  gay  suitors,  who  continue  performing 
strange  antics,  "until  a  marked  preference  is  shown  for  one." 
The  female  of  the  red-winged  starling  (Agelseus  phoeniceus)  Is 
likewise  pursued  by  several  males,  "until,  becoming  fatigued, 
"she  alights,  receives  their  addresses,  and  soon  makes  a  choice." 
He  describes  also  how  several  male  night-jars  repeatedly  plunge 
through  the  air  with  astonishing  rapidity,  suddenly  turning,  and 
thus  making  a  singular  noise,  "but  no  sooner  has  the  female 
"made  her  choice,  than  the  other  males  are  driven  away."  With 
one  of  the  vultures  (Cathartes  aura)  of  the  United  States,  parties 
of  eight,  ten,  or  more  males  and  females  assemble  on  fallen  logs. 


PREFERENCE  BY  THE  FEMALE.  411 

"exhibiting  the  strongest  desire  to  please  mutually,"  and  after 
many  caresses,  each  male  leads  off  his  partner  on  the  wing.  Audu- 
bon likewise  carefully  observed  the  wild  flocks  of  Canada  geese 
(.Anser  canadensis),  and  gives  a  graphic  description  of  their  love- 
antics;  he  says  that  the  birds  which  had  been  previously  mated 
"renewed  their  courtship  as  early  as  the  month  of  January,  while 
"the  others  would  be  contending  or  coquetting  for  hours  every  day, 
"until  all  seemed  satisfied  with  the  choice  they  had  made,  after 
"which,  although  they  remained  together,  any  person  could  easily 
"perceive  that  they  were  careful  to  keep  in  pairs.  I  have  ob- 
"served  also  that  the  older  the  birds,  the  shorter  were  the  pre- 
"liminaries  of  their  courtship.  The  bachelors  and  old  maids, 
"whether  in  regret,  or  not  earing  to  be  disturbed  by  the  bustle, 
"quietly  moved  aside  and  lay  down  at  some  distance  from  the 
"rest.''^**  Many  similar  statements  with  respect  to  other  birds 
could  be  cited  from  this  same  observer. 

Turning  now  to  domesticated  and  confined  birds,  I  will  com- 
mence by  giving  what  little  I  have  learnt  respecting  the  courtship 
of  fowls. 

I  have  received  long  letters  on  this  subject  from  Messrs.  Hewitt 
and  Tegetmeier,  and  almost  an  essay  from  the  late  Mr.  Brent.  It 
will  be  admitted  by  every  one  that  these  gentlemen,  so  well  known 
from  their  published  works,  are  careful  and  experienced  observers. 
They  do  not  believe  that  the  females  prefer  certain  males  on  ac- 
count of  the  beauty  of  their  plumage;  but  some  allowance  must  be 
made  for  the  artificial  state  under  which  these  birds  have  long  been 
kept.  Mr.  Tegetmeier  is  convinced  that  a  game-cock,  though  dis- 
figured by  being  dubbed  and  with  his  hackles  trimmed,  would  be 
accepted  as  readily  as  a  male  retaining  all  his  natural  ornaments. 
Mr.  Brent,  however,  admits  that  the  beauty  of  the  male  probably 
aids  in  exciting  the  female;  and  her  acquiescence  is  necessary. 
Mr.  Hewitt  is  convinced  that  the  union  is  by  no  means  left  to  mere 
chance,  for  the  female  almost  invariably  prefers  the  most  vigor- 
ous, defiant,  and  mettlesome  male;  hence  it  is  almost  useless,  as 
he  remarks,  "to  attempt  true  breeding  if  a  game-cock  in  good 
"health  and  condition  runs  the  locality,  for  almost  every  hen  on 
"leaving  the  roosting-place  will  resort  to  the  game-cock,  even 
"though  that  bird  may  not  actually  drive  away  the  male  of  her 
"own  variety."  Under  ordinary  circumstances  the  males  and  fe- 
males of  the  fowl  seem  to  come  to  a  mutual  understanding  by 
means  of  certain  gestures,  described  to  me  by  Mr.  Brent.  But 
hens  will  often  avoid  the  officious  attentions  of  young  males.  Old 
hens,  and  hens  of  a  pugnacious  disposition,  as  the  same  writer  in- 
forms me,  dislike  strange  males,  and  will  not  yield  until  well 

20  Audubon,  'Ornitholog.  Biography,'  vol  i.  pp.  191,  349;    vol.  il.  pp.  42, 
275;  vol.  iii.  p.  2. 


412  THE  DESCENT  OF  MAN. 

beaten  into  compliance.  Ferguson,  however,  describes  how  a 
quarrelsome  hen  was  subdued  by  the  gentle  courtship  of  a  Shang- 
hai cock.=^^ 

There  is  reason  to  believe  that  pigeons  of  both  sexes  prefer  pair- 
ing with  birds  of  the  same  breed;  and  dovecot-pigeons  dislike  all 
the  highly  improved  breeds.^  Mr.  Harrison  Weir  has  lately  heard 
from  a  trustworthy  observer,  who  keeps  blue  pigeons,  that  these 
drive  away  all  other  colored  varieties,  such  as  white,  red,  and  yel- 
low; and  from  another  observer,  that  a  female  dun  carrier  could 
not,  "after  repeated  trials,  be  matched  with  a  black  male,  but  im- 
mediately paired  with  a  dun.  Again.  Mr.  Tegetmeier  had  a  fe- 
male blue  turbit  that  obstinately  refused  to  pair  with  two  males  of 
the  same  breed,  which  were  successively  shut  up  with  her  for 
weeks;  but  on  being  let  out  she  would  have  immediately  accepted 
the  first  blue  dragon  that  offered.  As  she  was  a  valuable  bird,  she 
was  then  shut  up  for  many  weeks  with  a  silver  (i.e.,  very  pale  blue) 
male,  and  at  last  mated  with  him.  Nevertheless,  as  a  general  rule, 
color  appears  to  have  little  influence  on  the  pairing  of  pigeons. 
Mr.  Tegetmeier,  at  my  request,  stained  some  of  his  birds  with  ma- 
genta, but  they  were  not  much  noticed  by  the  others. 

Female  pigeons  occasionally  feel  a  strong  antipathy  towards  cer- 
tain males,  without  any  assignable  cause.  Thus  MM.  Boitard  and 
Corbie,  whose  experience  extended  over  forty-five  years,  state; 
"Quand  une  femelle  eprouve  de  I'antipathie  pour  un  male  avec 
"lequel  on  veut  Taccoupler,  malgre  tous  les  feux  de  I'amour, 
"malgre  I'alpiste  et  le  chenevis  dont  on  la  nourrit  pour  augmenter 
"son  ardeur,  malgre  un  emprisonnement  de  six  mois  et  meme  d'un 
"an,  elle  refuse  constamment  ses  caresses;  les  avances  empressees, 
"les  agaceries,  les  tournoiemens,  les  tendres  roucoulemens,  rien  ne 
"peut  lui  plaire  ni  I'emouvoir;  gonflee,  boudeuse,  blottie  dans  un 
"coin  de  sa  prison,  elle  n'en  sort  que  pour  boire  et  manger,  ou  pour 
"repousser  avec  une  espece  de  rage  des  caresses  devenues  trop 
"pressantes."^*  On  the  other  hand  Mr.  Harrison  Weir  has  himself 
observed,  and  has  heard  from  several  breeders,  that  a  female  pig- 
eon will  occasionally  take  a  strong  fancy  for  a  particular  male,  and 
will  desert  her  own  mate  for  him.  Some  females,  according  to 
another  experienced  observer,  Riedel,^*  are  of  a  profligate  dispo- 
sition, and  prefer  almost  any  stranger  to  their  own  mate.  Some 
amorous  males,  called  by  our  English  fanciers  "gay  birds,"  are 
so  successful  in  their  gallantries,  that,  as  Mr.  H.  Weir  informs  me, 

21  'Rare  and  Prize  Poultry,'  1854,  p.  27. 

22  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol. 
ii.  p.  103. 

28  Boitard  and  Corbie,  'Les  Pig-eons,  &c.,'  1824,  p.  12.  Prosper  Lucas 
(Traite  de  I'Hered.  Nat.'  torn,  ii,  1850,  p.  296)  has  himself  observed 
nearly  similar  facts  with  pigeons. 

»*  'Die  Taubenzucht,'  1824,  s.  86. 


PREB'ERENCE  BY  THE  FEMALE.  413 

they  must  be  shut  up  on  account  of  the  mischief  which  they 

cause. 

Wild  turkeys  in  the  United  States,  according  to  Audubon, 
"sometimes  pay  their  addresses  to  the  domesticated  females, 
"and  are  generally  received  by  them  with  great  pleasure."  So 
that  these  females  apparently  prefer  the  wild  to  their  own  males.=^-' 

Here  is  a  more  curious  case.  Sir  R.  Heron  during  many  years 
kept  an  account  of  the  habits  of  the  peafowl,  which  he  bred  in 
large  numbers.  He  states  that  "the  hens  have  frequently  great 
"preference  to  a  particular  peacock.  They  were  all  so  fond  of  an 
"old  pied  cock,  that  one  year,  when  he  was  confined  though  still 
"in  view,  they  were  constantly  assembled  close  to  the  trellice- 
"walls  of  the  prison,  and  would  not  suffer  a  japanned  peacock 
"to  touch  them.  On  his  being  let  out  in  the  autumn,  the  oldest  of 
"the  hens  instantly  courted  him,  and  was  successful  in  her  court- 
"ship.  The  next  year  he  was  shut  up  in  a  stable,  and  then  the 
"hens  all  courted  his  rival."^®  This  rival  was  a  japanned  or  black- 
winged  peacock,  to  our  eyes  a  more  beautiful  bird  than  the  com- 
mon kind. 

Lichtenstein,  who  was  a  good  observer  and  had  excellent  op- 
portunities of  observation  at  the  Cape  of  Good  Hope,  assured 
Rudolphi  that  the  female  widow-bird  (Chera  progne)  disowns  the 
male,  when  robbed  of  the  long  tail-feathers  with  which  he  is  orna- 
mented during  the  breeding-season.  I  presume  that  this  observa- 
tion must  have  been  made  on  birds  under  confinement."  Here  is 
an  analogous  case;  Dr.  Jaeger,-^  director  of  the  Zoological  Gar- 
dens, of  Vienna,  states  that  a  male  silver-pheasant,  who  had  been 
triumphant  over  all  other  males  and  was  the  accepted  lover  of  the 
females,  had  his  ornamental  plumage  spoiled.  He  was  then  im- 
mediately superseded  by  a  rival,  who  got  the  upper  hand  and  after- 
wards led  the  fiock. 

It  is  a  remarkable  fact,  as  showing  how  important  color  is  in 
the  courtship  of  birds,  that  Mr.  Boardman,  a  well-known  col- 
lector and  observer  of  birds  for  many  years  in  the  Northern 
United  States,  has  never  in  his  large  experience  seen  an  albino 
paired  with  another  bird;  yet  he  has  had  opportunities  of  ob- 
serving many  albinos  belonging  to  several  species.^    It  can  hardly 

25  'Ornithological  Biography,'  vol.  i.  p.  13.  See  to  the  same  effect. 
Dr.  Bryant,  in  'Allen's  Mammals  and  Birds  of  Florida,'  p.  344. 

26  «proc.  Zool.  Soc'  1835,  p.  54.  The  japanned  peacock  is  considered 
by  Mr.  Sclater  as  a  distinct  species,  and  has  been  named  Pavo  nigri- 
pennis;  but  the  evidence  seems  to  m©  to  show  that  it  is  only  a  variety. 

27  Rudolphi,  'Beytrage  zur  Anthropologie,'  1812,  s.  184. 

28  *Die  Darwin' sche  Theorie,  und  ihre  Stellung  zu  Moral  und  Re- 
ligion,' 1869,  s.  59. 

^»  This  statement  is  given  by  Mr.  A. .  Lelth  Adams,  in  his  'Field  and 
Forest  Rambles,'  1873,  p.  76,  and  accords  with  his  own  experience. 


414  THE  DESCENT  OP  MAN. 

be  maintained  that  albinos  in  a  state  of  nature  are  incapable  of 
breeding,  as  they  can  be  raised  with  the  greatest  facility  under 
confinement.  It  appears,  therefore,  that  we  must  attribute  the 
fact  that  they  do  not  pair,  to  their  rejection  by  their  normally  col- 
ored comrades. 

Female  birds  not  only  exert  a  choice,  but  in  some  few  cases  they 
court  the  male,  or  even  fight  together  for  his  possession.  Sir  R. 
Heron  states  that  with  peafowl,  the  first  advances  are  always 
made  by  the  female;  something  of  the  same  kind  takes  place,  ac- 
cording to  Audubon,  with  the  older  females  of  the  wild  turkey. 
With  the  capercailzie,  the  females  flit  round  the  male  whilst  he  is 
parading  at  one  of  the  places  of  assemblage,  and  solicit  his  at- 
tention.^" We  have  seen  that  a  tame  wild-duck  seduced  an  un- 
willing pintail  drake  after  a  long  courtship.  Mr.  Bartlett  believes 
that  the  Lophophorus,  like  many  other  gallinaceous  birds,  is  nat- 
urally polygamous,  but  two  females  cannot  be  placed  in  the  same 
cage  with  a  male,  as  they  fight  so  much  together.  The  following 
instance  of  rivalry  is  more  surprising  as  it  relates  to  bullfinches, 
which  usually  pair  for  life.  Mr.  Jenner  Weir  introduced  a  dull- 
colored  and  ugly  female  into  his  aviary,  and  she  immediately  at- 
tacked another  mated  female  so  unmercifully,  that  the  latter  had 
to  be  separated.  The  new  female  did  all  the  courtship,  and  was 
at  last  successful,  for  she  paired  with  the  male;  but  after  a  time 
she  met  with  a  just  retribution,  for,  ceasing  to  be  pugnacious,  she 
was  replaced  by  the  old  female,  and  the  male  then  deserted  his 
new  and  returned  to  his  old  love. 

In  all  ordinary  cases  the  male  is  so  eager  that  he  will  accept 
any  female,  and  does  not,  as  far  as  we  can  judge,  prefer  one  to  the 
other;  but,  as  we  shall  hereafter  see,  exceptions  to  this  rule  ap- 
parently occur  in  some  few  groups.  With  domesticated  birds,  I 
have  heard  of  only  one  case  of  males  showing  any  preference  for 
certain  females,  namely,  that  of  the  domestic  cock,  who,  accord- 
ing to  the  high  authority  of  Mr.  Hewitt,  prefers  the  younger  to  the 
older  hens.  On  the  other  hand,  in  effecting  hybrid  unions  between 
the  male  pheasant  and  common  hens,  Mr.  Hewitt  is  convinced 
that  the  pheasant  invariably  prefers  the  older  birds.  He  does  not 
appear  to  be  in  the  least  influenced  by  their  color,  but  "is  most 
"capricious  in  his  attachments;"^^  from  some  inexplicable  cause 
he  shows  the  most  determined  aversion  to  certain  hens,  which  no 
care  on  the  part  of  the  breeder  can  overcome.  Mr.  Hewitt  in- 
forms me  that  some  hens  are  quite  unattractive  even  to  the  males 

^  In  regard  to  peafowl,  see  Sir  R.  Heron,  'Proc.  Zoolog.  Soc'  1835,  p. 
54,  and  the  Rev.  E.  S.  Dixon,  'Ornamental  Poultry,'  1848,  p.  8.  For  the 
turkey,  Audubon,  ibid.  p.  4.  For  the  capercailzie,  Lloyd,  'Game  Birds 
of  Sweden,'  1867,  p.  23. 

31  Mr.  Hev/itt,  quoted  in  'Tegetmeier's  Poultry  Book,'  1866,  p.  165. 


PREFERENCE  BY  THE  FEMALE.  415 

of  their  own  species,  so  that  they  may  be  kept  with  several  cocks 
during  a  whole  season  and  not  one  egg  out  of  forty  or  fifty  will 
prove  fertile.  On  the  other  hand,  with  the  Long-tailed  duck 
(Harelda  glacialis),  "it  has  been  remarked,"  says  M.  Ekstrom, 
"that  certain  females  are  much  more  courted  than  the  rest.  Fre- 
'•'quently,  indeed,  one  sees  an  individual  surrounded  by  six  or 
"eight  amorous  males."  Whether  this  statement  is  credible,  I 
know  not;  but  the  native  sportsman  shoot  these  females  in  order 
to  stuf£  them  as  decoys.^- 

With  respect  to  female  birds  feeling  a  preference  for  particular 
males,  we  must  bear  in  mind  that  we  can  judge  of  choice  being 
exerted,  only  by  analogy.  If  an  inhabitant  of  another  planet  were 
to  behold  a  number  of  young  rustics  at  a  fair  courting  a  pretty 
girl,  and  quarreling  about  her  like  birds  at  one  of  their  places  of 
assemblage,  he  would,  by  the  eagerness  of  the  wooers  to  please  her 
and  to  display  their  finery,  infer  that  she  had  the  power  of  choice. 
Now  with  birds,  the  evidence  stands  thus;  they  have  acute  powers 
of  observation,  and  they  seem  to  have  some  taste  for  the  beautiful 
both  in  color  and  sound.  It  is  certain  that  the  females  occa- 
sionally exhibit,  from  unknown  causes,  the  strongest  antipathies 
and  preferences  for  particular  males.  When  the  sexes  differ  in 
color  or  in  other  ornaments  the  males  with  rare  exceptions  are 
the  more  decorated,  either  permanently  or  temporarily  during 
the  breeding-season.  They  sedulously  display  their  various  or- 
naments, exert  their  voices,  and  perform  strange  antics  in  the 
presence  of  the  females.  Even  well-armed  males,  who,  it  might 
be  thought,  would  altogether  depend  for  success  on  the  law  of  bat- 
tle, are  in  most  cases  highly  ornamented;  and  their  ornaments 
have  been  acquired  at  the  expense  of  some  loss  of  power.  In  other 
cases  ornaments  have  been  acquired,  at  the  cost  of  increased  risk 
from  birds  and  beasts  of  prey.  With  various  species  many  indi- 
viduals of  both  sexes  congregate  at  the  same  spot,  and  their  court- 
ship is  a  prolonged  affair.  There  is  even  reason  to  suspect  that 
the  males  and  females  within  the  same  district  do  not  always 
succeed  in  pleasing  each  other  and  pairing. 

What  then  are  we  to  conclude  from  these  facts  and  considera- 
tions? Does  the  male  parade  his  charms  with  so  much  pomp  and 
rivalry  for  no  purpose?  Are  we  not  justified  in  believing  that  the 
female  exerts  a  choice,  and  that  she  receives  the  addresses  of  the 
male  who  pleases  her  most?  It  is  not  probable  that  she  con- 
sciously deliberates;  but  3he  is  most  excited  or  attracted  by  the 
most  beautiful,  or  melodious,  or  gallant  males.  Nor  need  it  be 
supposed  that  the  female  studies  each  stripe  or  spot  of  color;  that 
the  peahen,  for  instance,  admires  each  detail  in  the  gorgeous 
train  of  the  peacock — she  is  probably  struck  only  by  the  general 


Quoted  in  Lloyd's  'Game  Birds  of  Sweden,'  p.  345. 


416  THE  DESCENT  OF  MAN. 

effect.  Nevertheless,  after  hearing  how  carefully  the  male  Argus 
pheasant  displays  his  elegant  primary  wing-feathers,  and  erects 
his  ocellated  plumes  in  the  right  position  for  their  full  effect;  or 
again,  how  the  male  goldfinch  alternately  displays  his  gold-be- 
spangled wings,  we  ought  not  to  feel  too  sure  that  the  female  does 
not  attend  to  each  detail  of  beauty.  We  can  judge,  as  already  re- 
marked, of  choice  being  exerted,  only  from  analogy;  and  the  men- 
tal powers  of  birds  do  not  differ  fundamentally  from  ours.  From 
these  various  considerations  we  may  conclude  that  the  pairing  of 
birds  is  not  left  to  chance;  but  that  those  males,  which  are  best 
able  by  their  various  charms  to  please  or  excite  the-  female,  are 
under  ordinary  circumstances  accepted.  If  this  be  admitted,  there 
is  not  much  difficulty  in  understanding  how  male  birds  have  grad- 
ually acquired  their  ornamental  characters.  All  animals  present 
individual  differences,  and  as  man  can  modify  his  domesticated 
birds  by  selecting  the  individuals  which  appear  to  him  the  most 
beautiful,  so  the  habitual  or  even  occasional  preference  by  the 
female  of  the  more  attractive  males  would  almost  certainly  lead 
to  their  modification;  and  such  modifications  might  in  the  course 
of  time  be  augmented  to  almost  any  extent,  compatible  with  the 
existence  of  the  species. 

Variability  of  Birds,  and  especially  of  their  Secondary  Sexual 
Characters. — Variability  and  inheritance  are  the  foundations  for 
the  work  of  selection.  That  domesticated  birds  have  varied  great- 
ly, their  variations  being  inherited,  is  certain.  That  birds  in  a 
state  of  nature  have  been  modified  into  distinct  races  is  now 
universally  admitted.^^  Variations  may  be  divided  into  two 
classes;  those  which  appear  to  our  ignorance  to  arise  spontane- 
ously, and  those  which  are  directly  related  to  the  surrounding 
conditions,  so  that  all  or  nearly  all  the  individuals  of  the  same 
species  are  similarly  modified.  Cases  of  the  latter  kind  have 
recently  been  observed  with  care  by  Mr.  J.  A.  Allen,^*  who  shows 


33  According  to  Dr.  Blasius  (Tbis,'  vol.  ii.  1860,  p.  297),  there  are  425 
indubitable  species  of  birds  which  breed  in  Europe,  besides  sixty  forms, 
which  are  frequently  regarded  as  distinct  species.  Of  the  latter, 
Blasius  thinks  that  only  ten  are  really  doubtful,  and  that  the  other  fifty 
ought  to  be  united  with  their  nearest  allies;  but  this  shows  that  there 
must  be  a  considerable  amount  of  variation  with  some  of  our  Euro- 
pean birds.  It  is  also  an  unsettled  point  with  naturalists,  whether 
several  North  American  birds  ought  to  be  ranked  as  specifically  dis- 
tinct from  the  corresponding  European  species.  So  again  many  North 
American  forms  which  until  lately  were  named  as  distinct  species, 
are  now  considered  to  be  local  races. 

^  'Mammals  and  Birds  of  East  Florida,'  also  an  'Ornithological 
Reconnoissance  of  Kansas,'  &c.  Notwithstanding  the  influence  of 
climate  on  the  colors  of  birds,  it  is  difficult  to  account  for  the  dull 
or  dark  tints  of  almost  all  the  species  inhabiting  certain  countries. 


VARIABILITY.  417 

that  in  the  United  States  many  species  of  birds  gradually  become 
more  strongly  colored  in  proceeding  southward,  and  more  lightly 
colored  in  proceeding  westward  to  the  arid  plains  of  the  interior. 
Both  sexes  seem  generally  to  be  affected  in  a  like  manner,  but 
sometimes  one  sex  more  than  the  other.  This  result  is  not  in- 
compatible with  the  belief  that  the  colors  of  birds  are  mainly 
due  to  the  accumulation  of  successive  variations  through  sexual 
selection;  for  even  after  the  sexes  have  been  greatly  differen- 
tiated, climate  might  produce  an  equal  effect  on  both  sexes,  or  a 
greater  effect  on  one  sex  than  on  the  other,  owing  to  some  con* 
stitutional  difference. 

Individual  differences  between  the  members  of  the  same  spe- 
cies are  admitted  by  every  one  to  occur  under  a  state  of  nature. 
Sudden  and  strongly  marked  variations  are  rare;  it  is  also  doubt- 
ful whether  if  beneficial  they  would  often  be  preserved  through 
selection  and  transmitted  to  succeeding  generations.^"  Neverthe- 
less, it  may  be  worth  while  to  give  the  few  cases  which  I  have  been 
able  to  collect,  relating  chiefly  to  color, — simple  albinism  and 
melanism  being  excluded.  Mr.  Gould  is  well  known  to  admit 
the  existence  of  a  few  varieties,  for  he  esteems  very  slight  differ- 
ences as  specific;  yet  he  states^*^  that  near  Bogota  certain  hum- 
ming-birds belonging  to  the  genus  Cynanthus  are  divided  into 

for  instance,  the  Galapagos  Islands  under  the  equator,  the  wide  tem- 
perate plains  of  Patagonia,  and,  as  it  appears,  Egypt  (see  Mr. 
Hartshorne  in  the  'American  Naturalist,'  1873,  p.  747).  These  coun- 
tries are  open,  and  afford  little  shelter  to  birds;  but  it  seems  doubtful 
whether  the  absence  of  brightly  colored  species  can  he  explained  on  the 
principle  of  protection,  for  on  the  Pampas,  which  are  equally  open, 
though  covered  by  green  grass,  and  where  the  birds  would  be  equally 
exposed  to  danger,  many  brilliant  and  conspicuously  colored  species 
are  common.  I  have  sometimes  speculated  whether  the  prevailing 
dull  tints  of  the  scenery  in  the  above  named  countries  may  not  have 
affected  the  appreciation  of  bright  colors  by  the  birds  inhabiting  them. 

35  'Origin  of  Species,'  fifth  edit.  1869,  p.  104.  I  had  always  perceived, 
that  rare  and  strongly-marked  deviations  of  structure,  deserving  to 
bo  called  monstrosities,  could  seldom  be  preserved  through  natural 
selection,  and  that  the  preservation  of  even  highly-beneficial  varia- 
tions would  depend  to  a  certain  extent  on  chance.  I  had  also  fully 
appreciated  the  importance  of  mere  individual  differences,  and  this 
led  me  to  insist  so  strongly  on  the  importance  of  that  unconscious 
form  of  selection  by  man,  which  follows  from  the  preservation  of  the 
most  valued  individuals  of  each  breed,  without  any  intention  on  Kis 
part  to  modify  the  characters  of  the  breed.  But  until  I  read  an  able 
article  in  the  'North  British  Review'  (March,  1867,  p.  289,  et  seq.), 
which  has  been  of  more  use  to  me  than  any  other  Review,  I  did  not 
see  how  great  the  chances  were  against  the  preservation  of  variations, 
whether  slight  or  strongly  pronounced,  occurring  only  in  single  indi- 
viduals. 

36  'introduct.  to  the  Trochilidae,'  p.  102. 

2$ 


418  THE  DESCENT  OF  MAN. 

two  or  three  races  or  varieties,  which  differ  from  each  other  in 
the  coloring  of  the  tail — "some  having  the  whole  of  the  feathers 
"blue,  while  others  have  the  eight  central  ones  tipped  with  beau- 
"tiful  green."  It  does  not  appear  that  intermediate  gradations 
have  been  observed  in  this  or  the  following  cases.  In  the  males 
alone  of  one  of  the  Australian  parrakeets,  "the  thighs  in  some  are 
"scarlet,  in  others  grass-green."  In  another  parrakeet  of  the 
same  country  "some  individuals  have  the  band  across  the  wing- 
"coverts  bright-yellow,  while  in  others  the  same  part  is  tinged 
"with  red.""  In  the  United  States  some  few  of  the  males  of  the 
Scarlet  Tanager  (Tanagra  rubra)  have  "a  beautiful  transverse 
"band  of  glowing  red  on  the  smaller  v/ing-coverts;"^**  but  this 
variation  seems  to  be  somewhat  rare,  so  that  its  preservation 
through  sexual  selection  would  follow  only  under  unusually  favor- 
able circumstances.  In  Bengal  the  Honey  buzzard  (Pernis  cris- 
tata)  has  either  a  small  rudimental  crest  on  its  head,  or  none  at 
all:  so  slight  a  difference,  however,  would  not  have  been  worth 
notice,  had  not  this  same  species  possessed  in  Southern  India  "a 
"well-marked  occipital  crest  formed  of  several  graduated  feath- 


"39 


"ers. 

The  following  case  is  in  some  respects  more  interesting.  A 
pied  variety  of  the  raven,  with  the  head,  breast,  abdomen,  and 
parts  of  the  wings  and  tail-feathers  white,  is  confined  to  the 
Feroe  Islands.  It  is  not  very  rare  there,  for  Graba  saw  during 
his  visit  from  eight  to  ten  living  specimens.  Although  the  char- 
acters of  this  variety  are  not  quite  constant,  yet  it  has  been  named 
by  several  distinguished  ornithologists  as  a  distinct  species.  The 
fact  of  the  pied  birds  being  pursued  and  persecuted  with  much 
clamor  by  the  other  ravens  of  the  island  was  the  chief  cause 
which  led  Briinnich  to  conclude  that  they  were  specifically  dis- 
tinct; but  this  is  now  known  to  be  an  error.*"  This  case  seems 
analogous  to  that  lately  given  of  albino  birds  not  pairing  from 
being  rejected  by  their  comrades. 

In  various  parts  of  the  northern  seas  a  remarkable  variety  of 
the  common  Guillemot  (Uria  troile)  is  found;  and  in  Feroe  one 
out  of  every  five  birds,  according  to  Graba's  estimation,  presents 
this  variation.  It  is  characterized*^  by  a  pure  white  ring  round 
the  eye,  with  a  curved  narrow  white  line,  an  inch  and  a  half  in 
length,  extending  back  from  the  ring.    This  conspicuous  charac- 

37  Gould,  'Handbook  to  Birds  of  Australia/  vol.  ii.  pp.  32  and  68. 

38  Audubon,  'Ornitholog.  Biography,'  1838,  vol.  iv.  p.  389. 

3»  Jerdon,  'Birds  of  India,'  vol.  i.  p.  108;  and  Mr.  Blyth,  in  'Land  and 
Water,'  1868,  p.  381. 

40  Graba,  'Tagebuch,  Relse  nach  Faro,'  1830,  s.  51-54,  Macgillivray, 
'Hist.  British  Birds,'  vol.  iii.  p.  745.    'Ibis,'  vol.  v.  1863,  p.  469. 

^  Graba,  ibid.  s.  54.    Macgillivray,  ibid.  vol.  v.  p.  327. 


VARIABILITY.  419 

ter  has  caused  the  bird  to  be  ranked  by  several  ornithologists 
as  a  distinct  species  under  the  name  of  U.  lacrymans,  but  it  is 
now  known  to  be  merely  a  variety.  It  often  pairs  with  the  com- 
mon kind,  yet  intermediate  gradations  have  never  been  seen;  nor 
is  this  surprising,  for  variations  which  appear  suddenly,  are  often, 
as  I  have  elsewhere  shown,*-  transmitted  either  unaltered  or  not 
at  all.  We  thus  see  that  two  distinct  forms  of  the  same  species 
may  co-exist  in  the  same  district,  and  we  cannot  doubt  that  if 
the  one  had  possessed  any  advantage  over  the  other,  it  would 
soon  have  been  multiplied  to  the  exclusion  of  the  latter.  If,  for 
instance,  the  male  pied  ravens,  instead  of  being  persecuted  by 
their  comrades,  had  been  highly  attractive  (like  the  above  pied 
peacock)  to  the  black  female  ravens,  their  numbers  would  have 
rapidly  increased.  And  this  would  have  been  a  case  of  sexual 
selection. 

With  respect  to  the  slight  individual  differences  which  are 
common,  in  a  greater  or  less  degree,  to  all  the  members  of  the 
same  species,  we  have  every  reason  to  believe  that  they  are  by 
far  the  most  important  for  the  work  of  selection.  Secondary 
sexual  characters  are  eminently  liable  to  varj'',  both  with  animals 
in  a  state  of  nature  and  under  domestication.*'  There  is  also 
reason  to  believe,  as  we  have  seen  in  our  eighth  chapter,  that 
variations  are  more  apt  to  occur  in  the  male  than  in  the  female 
sex.  All  these  contingencies  are  highly  favorable  for  sexual 
selection.  Whether  characters  thus  acquired  are  transmitted  to 
one  sex,  or  to  both  sexes,  depends,  as  we  shall  see  in  the  fol- 
lowing chapter,  on  the  form  of  inheritance  which  prevails. 

It  is  sometimes  difficult  to  form  an  opinion  whether  certain 
slight  differences  between  the  sexes  of  birds  are  simply  the  result 
of  variability  with  sexually-limited  inheritance,  without  the  aid 
of  sexual  selection,  or  whether  they  have  been  augmented  through 
this  latter  process.  I  do  not  here  refer  to  the  many  instances 
where  the  male  displays  splendid  colors  or  other  ornaments,  of 
which  the  female  partakes  to  a  slight  degree;  for  these  are  al- 
most certainly  due  to  characters  primarily  acquired  by  the  male 
having  been  more  or  less  transferred  to  the  female.  But  what  are 
we  to  conclude  with  respect  to  certain  birds  in  which,  for  instance, 
the  eyes  differ  slightly  in  color  in  the  two  sexes?**  In  some  cases 
the  eyes  differ  conspicuously;  thus  with  the  storks  of  the  genus 
Xenorhynchus,  those  of  the  male  are  blackish-hazel,  whilst  those 


42  'Tariation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
p.  92. 

*3  On  these  points  see,  also,  'Variation  of  Animals  and  Plants  under 
Domestication,'  vol.  i.  p.  253;  vol.  ii.  pp.  73,  75. 

**  See,  for  instance,  on  the  irides  of  a  Podica  and  Gallicrex  in  'Ibis," 
vol,  ii.  1860,  p.  206;    and  vol.  v.  1863,  p.  426. 


420  THE  DESCENT  OF  MAN. 

of  the  female  are  gamboge-yellow;  with  many  hornbills  (Bii- 
ceros),  as  I  hear  from  Mr.  Blyth,^^  the  males  have  intense  crimsoa 
eyes,  and  those  of  the  females  are  white.  In  the  Buceros  bicornis, 
the  hind  margin  of  the  casque  and  a  stripe  on  the  crest  of  the 
beak  are  black  in  the  male,  but  not  so  in  the  female.  Are  we  to 
suppose  that  these  black  marks  and  the  crimson  color  of  the  eyes 
have  been  preserved  or  augmented  through  sexual  selection  in 
the  males?  This  is  very  doubtful;  for  Mr.  Bartlett  showed  me  m 
the  Zoological  G-ardens  that  the  inside  of  the  mouth  of  this 
Buceros  is  black  in  the  male  and  flesh-colored  in  the  female;  and 
their  external  appearance  or  beauty  would  not  be  thus  affected. 
I  observed  in  Chili*^  that  the  iris  in  the  condor,  when  about  a 
year  old,  is  dark-brown,  but  changes  at  maturity  into  yellowish- 
brown  in  the  male,  and  into  bright  red  in  the  female.  The  male 
has  also  a  small,  longitudinal,  leaden-colored,  fleshy  crest  or 
comb.  The  comb  of  many  gallinaceous  birds  is  highly  ornament- 
al, and  assumes  vivid  colors  during  the  act  of  courtship;  but 
what  are  we  to  think  of  the  dull-colored  comb  of  the  condor, 
which  does  not  appear  to  us  in  the  least  ornamental?  The  same 
question  may  be  asked  in  regard  to  various  other  characters, 
such  as  the  knob  on  the  base  of  the  beak  of  the  Chinese  goose 
(Anser  cygnoides),  which  is  much  larger  in  the  male  than  in  the 
female.  No  certain  answer  can  be  given  to  these  questions;  but 
we  ought  to  be  cautious  in  assuming  that  knobs  and  various  flesby 
appendages  cannot  be  attractive  to  the  female,  when  we  remember 
that  with  savage  races  of  man  various  hideous  deformities — deep 
scars  on  the  face  with  the  flesh  raised  into  protuberances,  tiie 
septum  of  the  nose  pierced  by  sticks  or  bones,  holes  in  the  ears 
and  lips  stretched  widely  open — are  all  admired  as  ornamental. 

Whether  or  not  unimportant  differences  between  the  sexes, 
such  as  those  just  specified,  have  been  preserved  through  sexual 
selection,  these  differences,  as  well  as  all  others,  must  primarily 
depend  on  the  laws  of  variation.  On  the  principle  of  correlated 
development,  the  plumage  often  varies  on  different  parts  of  the 
body,  or  over  the  whole  body,  in  the  same  manner.  We  see  this 
well  illustrated  in  certain  breeds  of  the  fowl.  In  all  the  breeds 
the  feathers  on  the  neck  and  loins  of  the  males  are  elongated, 
and  are  called  hackles;  now  when  both  sexes  acquire  a  top-knot, 
which  is  a  new  character  in  the  genus,  the  feathers  on  the  head 
of  the  male  become  hackle-shaped,  evidently  on  the  principle  of 
correlation;  whilst  those  on  the  head  of  the  female  are  of  the 
ordinary  shape.  The  color  also  of  the  hackles  forming  the  top- 
knot of  the  male,  is  often  correlated  with  that  of  the  hackles  on 
the  neck  and  loins,  as  may  be  seen  by  comparing  these  feathers 

^  See,  also,  Jerdoii,   'Birds  of  India,'  vol.  i.  pp.  243-245. 
«  'Zoology  of  the  Voyage  of  H.  M.  S.  Beagle,'  1841,  p.  6. 


OCELLI.  421 

!n  the  Golden  and  Silver-spangled  Polisli,  the  Houdans,  and 
Creve-cceur  breeds.  In  some  natural  species  we  may  ohserve 
exactly  the  same  correlation  in  the  colors  of  these  same  feathers, 
as  in  the  males  of  the  splendid  Gold  and  Amherst  pheasants. 

The  structure  of  each  individual  feather  generally  causes  any 
change  in  its  coloring  to  be  symmetrical;  we  see  this  in  the 
various  laced,  spangled,  and  penciled  breeds  of  the  fowl;  and 
on  the  principle  of  correlation  the  feathers  over  the  whole  body 
are  often  colored  in  the  same  manner.  We  are  thus  enabled 
without  much  trouble  to  rear  breeds  with  their  plumage  marked 
almost  as  symmetrically  as  in  natural  species.  In  laced  and 
spangled  fowls  the  colored  margins  of  the  feathers  are  abruptly 
defined;  but  in  a  mongrel  raised  by  me  from  a  black  Spanish 
cock  glossed  with  green,  and  a  white  game-hen,  all  the  feathers 
were  greenish-black,  excepting  towards  their  extremities,  which 
were  yellowish-white;  but  between  the  white  extremities  and  the 
black  bases,  there  was  on  each  feather  a  symmetrical,  curved 
zone  of  dark-brown.  In  some  instances  the  shaft  of  the  feather 
determines  the  distribution  of  the  tints;  thus  with  the  body- 
feathers  of  a  mongrel  from  the  same  black  Spanish  cock  and  a 
silver-spangled  Polish  hen,  the  shaft,  together  with  a  narrow 
space  on  each  side,  was  greenish-black,  and  this  was  surrounded 
by  a  regular  zone  of  dark-brown,  edged  with  brownish-white.  In 
these  cases  we  have  feathers  symmetrically  shaded,  like  those 
which  give  so  much  elegance  to  the  plumage  of  many  natural 
species.  I  have  also  noticed  a  variety  of  the  common  pigeon 
with  the  wing-bars  symmetrically  zoned  with  three  bright  shades, 
instead  of  being  simply  black  on  a  slaty-blue  ground,  as  in  the 
parent-species. 

In  many  groups  of  birds  the  plumage  is  differently  colored 
in  the  several  species,  yet  certain  spots,  marks,  or  stripes  are 
retained  by  all.  Analogous  cases  occur  with  the  breeds  of  the 
pigeon,  v/hich  usually  retain  the  two  wing-bars,  though  they 
may  be  colored  red,  yellow,  white,  black,  or  blue,  the  rest  of  the 
plumage  being  of  some  wholly  different  tint.  Here  is  a  more 
curious  case,  in  which  certain  marks  are  retained,  though  col- 
ored in  a  manner  almost  exactly  the  opposite  of  what  is  natural; 
the  aboriginal  pigeon  has  a  blue  tail,  with  the  terminal  halves 
of  the  outer  webs  of  the  two  outer  tail  feathers  white;  now  there 
is  a  sub-variety  having  a  white  instead  of  a  blue  tail,  with  precise- 
ly that  part  black  which  is  white  in  the  parent  species.*^ 

Formation  and  Variability  of  the  Ocelli  or  eye-like  Spots  on  the 
Plumage  of  Birds.  — As  no  ornaments  are  more  beautiful  than  the 

*'  Bechstein,   'Naturg-eschichte  Deutscblands,'  B,  iv.  1735,  s.   31,  on  a 
STib-variety  of  the  Monck  pigeon. 
23 


422  THE  DESCENT  OF  MAN. 

ocelli  on  th*  feathers  of  various  birds,  on  the  hairy  coats  of  some 
mammals,  on  the  scales  of  reptiles  and  fishes,  on  the  skin  of 
amphibians,  on  the  wings  of  many  Lepidoptera  and  other  insects, 
they  deserve  to  be  especially  noticed.  An  ocellus  consists  of  a 
spot  within  a  ring  of  another  color,  like  the  pupil  within  the 
iris,  but  the  central  spot  is  often  surrounded  by  additional  con- 
centric zones.  The  ocelli  on  the  tail-coverts  of  the  peacock  offer 
a  familiar  example,  as  well  as  those  on  the  wings  of  the  peacock- 
butterfly  (Vanessa).  Mr.  Trimen  has  given  me  a  description  of 
a  S.  African  moth  (Gyananis  isis),  allied  to  our  Emperor  moth, 
in  which  a  magnificent  ocellus  occupies  nearly  the  whole  surface 
of  each  hinder  wing;  it  consists  of  a  black  center,  including  a 
semi-transparent  crescent-shaped  mark,  surrounded  by  successive, 
ochre-yellow,  black,  ochre-yellow,  pink,  white,  pink,  brown,  and 
whitish  zones.  Although  we  do  not  know  the  steps  by  which 
these  wonderfully  beautiful  and  complex  ornaments  have  been 
developed,  the  process  has  probably  been  a  simple  one,  at  least 
with  insects;  for,  as  Mr.  Trimen  writes  to  me,  "no  characters  of 
"mere  marking  or  coloration  are  so  unstable  in  the  Lepidoptera 
"as  the  ocelli,  both  in  number  and  size."  Mr.  Wallace,  who  first 
called  my  attention  to  this  subject,  showed  me  a  series  of  speci- 
mens of  our  common  meadow-brown  butterfly  (Hipparchia  janira) 
exhibiting  numerous  gradations  from  a  simple  minute  black 
spot  to  an  elegantly-shaded  ocellus.  In  a  S.  African  butterfly 
(Cyllo  leda,  Linn.),  belonging  to  the  same  family,  the  ocelli  are 
even  still  more  variable.  In  some  specimens  (A,  fig.  53)  large 
spaces  on  the  upper  surface  of  the  wings  are  colored  black,  and 
include  irregular  white  marks;  and  from  this  state  a  complete 
gradation  can  be  traced  into  a  tolerably  perfect  ocellus  (A"), 
and  this  results  from  the  contraction  of  the  irregular  blotches 
of  color.  In  another  series  of  specimens  a  gradation  can  be  fol- 
lowed from  excessively  minute  white  dots,  surrounded  by  a  scarce- 
ly visible  black  line  (B),  into  perfectly  symmetrical  and  large 
ocelli  (B).^  In  cases  like  these  the  development  of  a  perfect 
ocellus  does  not  require  a  long  course  of  variation  and  selection. 
With  birds  and  many  other  animals,  it  seems  to  follow  from 
the  comparison  of  allied  species  that  circular  spots  are  often 
generated  by  the  breaking  up  and  contraction  of  stripes.  In  the 
Tragopan  pheasant  faint  white  lines  in  the  female  represent  the 
beautiful  white  spots  in  the  male;^  and  something  of  the  same 
kind  may  be  observed  in  the  two  sexes  of  the  Argus  pheasant. 


*8  This  woodcut  has  been  engraved  from  a  beautiful  drawing-,  most 
kindly  made  for  me  by  Mr.  Trimen;  see,  also,  his  description  of  the 
wonderful  amount  of  variation  in  the  coloration  and  shape  of  the 
wings  of  this  butterfly,  in  his  'Rhopalocera  Africae  Australis,'  p.  186. 

*»  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  517. 


OCELLI. 


423 


However  this  may  be,  appearances  strongly  favor  the  belief  that 
on  the  one  hand,  a  dark  spot  is  often  formed  by  the  coloring 
matter  being  drawn  towards  a  central  point  from  a  surrounding 
zone,  which  latter  is  thus  rendered  lighter;  and,  on  the  other 
hand,  that  a  white  spot  is  often  formed  by  the  color  being  driven 
away  from  a  central  point,  so  that  it  accumulates  in  a  surround- 
ing darker  zone.    In  either  case  an  ocellus  is  the  result.    The 


Fig.   53.    Cyllo  leda,  Linn.,   from  a  drawing  by  Mr.   Trimen,   showing 
the  extreme  range  of  variation  in  the  ocelli. 


A.  Specimen,  from  Mauritius,  up- 
per surface  of  fore-wing. 
A^.  Specimen,  from  Natal,  ditto. 


B.  Specimen,  from  Java,  upper 
surface  of  hind- wing. 

B^.  Specimen,  from  Mauritius, 
ditto. 


coloring  matter  seems  to  be  a  nearly  constant  quantity,  but  is 
redistributed,  either  centripetally  or  centrifugally.  The  feathers 
of  the  common  guinea-fowl  offer  a  good  instance  of  white  spots 
surrounded  by  darker  zones;  and  wherever  the  white  spots  are 
large  and  stand  near  each  other,  the  surrounding  dark  zones 
become  confluent.  In  the  same  wing-feather  of  the  Argus  pheas- 
ant dark  spots  may  be  seen  surrounded  by  a  pale  zone,  and  white 
spots  by  a  dark  zone.  Thus  the  formation  of  an  ocellus  in  its 
most  elementary  state  appears  to  be  a  simple  affair.  By  what 
further  steps  the  more  complex  ocelli,  which  are  surrounded  by 
many  successive  zones  of  color,  have  been  generated,  I  will  not 
pretend  to  say.  But  the  zoned  feathers  of  the  mongrels  from  dif- 
ferently colored  fowls,  and  the  extraordinary  variability  of  the 


424  THE  DESCENT  OF  MAN. 

ocelli  on  many  Lepidoptera,  lead  us  to  conclude  that  their  forma- 
tion is  not  a  complex  process,  but  depends  on  some  slight  and 
graduated  change  in  the  nature  of  the  adjoining  tissues. 

Oradation  of  Secondary  Sexual  Characters.  ■ — Cases  of  gradation 
are  important,  as  showing  us  that  highly  complex  ornaments 
may  be  acquired  by  small  successive  steps.  In  order  to  discover 
the  actual  steps  by  which  the  male  of  any  existing  bird  has 
acquired  his  magnificent  colors  or  other  ornaments,  we  ought 
to  behold  the  long  line  of  his  extinct  progenitors;  but  this  is 
obviously  impossible.  We  may,  however,  generally  gain  a  clue 
by  comparing  all  the  species  of  the  same  group,  if  it  be  a  large 
one;  for  some  of  them  will  probably  retain,  at  least  partially, 
traces  of  their  former  characters.  Instead  of  entering  on  tedious 
details  respecting  various  groups,  in  which  striking  instances 
of  gradation  could  be  given,  it  seems  the  best  plan  to  take  one 
or  two  strongly  marked  cases,  for  instance  that  of  the  peacock, 
in  order  to  see  if  light  can  be  thrown  on  the  steps  by  which 
this  bird  has  become  so  splendidly  decorated.  The  peacock  is 
chiefly  remarkable  from  the  extraordinary  length  of  his  tail- 
coverts;  the  tail  itself  not  being  much  elongated.  The  barbs 
along  nearly  the  whole  length  of  these  feathers  stand  separate  or 
are  decomposed;  but  this  is  the  case  with  the  feathers  of  many 
species,  and  with  some  varieties  of  the  domestic  fowl  and  pigeon. 
The  barbs  coalesce  towards  the  extremity  of  the  shaft  forming 
the  oval  disc  or  ocellus,  which  is  certainly  one  of  the  most  beau- 
tiful objects  in  the  world.  It  consists  of  an  iridescent,  intensely 
blue,  indented  center,  surrounded  by  a  rich  green  zone,  this  by  a 
broad  coppery-brown  zone,  and  this  by  five  other  narrow  zones 
of  slightly  different  iridescent  shades.  A  trifling  character  in 
the  disc  deserves  notice;  the  barbs,  for  a  space  along  one  ol 
the  concentric  zones  are  more  or  less  destitute  of  their  barbules, 
so  that  a  part  of  the  disc  is  surrounded  by  an  almost  transparent 
zone,  which  gives  it  a  highly  finished  aspect.  But  I  have  else- 
where described^°  an  exactly  analogous  variation  in  the  hackles 
of  a  sub-variety  of  the  game-cock,  in  which  the  tips,  having  a 
metallic  luster,  "are  separated  from  the  lower  part  of  the  feather 
"by  a  symmetrically  shaped  transparent  zone,  composed  of  the 
"naked  portions  of  the  barbs."  The  lower  margin  or  base  of  the 
dark-blue  center  of  the  ocellus  is  deeply  indented  on  the  line  of 
the  shaft.  The  surrounding  zones  likewise  show  traces,  as  may 
be  seen  in  the  drawing  (fig.  54),  of  indentations,  or  rather  breaks. 
These  indentations  are  common  to  the  Indian  and  Javan  peacocks 
(Pavo  cristatus  and  P.  muticus) ;  and  they  seemed  to  deserve  par- 
ticular attention,  as  probably  connected  with  the  development  of 


«°  'Variation  of  Animals  and  Plants  under  Domestication,'  vol.  i.  p.  254, 


GRADATION  OF  CHARACTERS. 


42.: 


the   ocellus;     but  for  a  long  time  I  could  not  conjecture  their 
meaning. 

If  we  admit  the  principle  of  gradual  evolution,  there  must 
formerly  have  existed  many  species  which  presented  every  suc- 
cessive step  between  the  wonderfully  elongated  tail-coverts  of 
the  peacock  and  the  short  tail-coverts  of  all  ordinary  birds;  and 
again  between  the  magnificent  ocelli  of  the  former,  and  the  sim- 
pler ocelli  or  mere  colored  spots  on  other  birds;  and  so  with  all 
the  other  characters  of  the  peacock.     Let  us  look  to  the  allied 


Fig.  54.  Feather  of  Peacock,  about  two-thirds  of  natural  size,  drawn 
by  Mr.  Ford.  The  transparent  zone  is  represented  by  the  outermost 
white  zone,  confined  to  the  upper  end  of  the  disc. 

Gallinacese  for  any  still-existing  gradations.  The  species  and 
sub-species  of  Polyplectron  inhabit  countries  adjacent  to  the 
native  land  of  the  peacock;  and  they  so  far  resemble  this  bird 
that  they  are  sometimes  called  peacock-pheasants.  I  am  also  in- 
formed by  Mr.  Bartlett  that  they  resemble  the  peacock  in  their 
voice  and  in  some  of  their  habits.  During  the  spring  the  males, 
as  previously  described,  strut  about  before  the  comparatively 
plain-colored  females,  expanding  and  erecting  their  tail  and  wing- 


426  THE  DESCENT  OF  MAN. 

feathers,  which  are  ornamented  with  numerous  ocelli.  I  re- 
quest the  reader  to  turn  back  to  the  drawing  (fig.  51,  p.  391)  of  a 
Polyplectron.  In  P.  napoleonis  the  ocelli  are  confined  to  the 
tail  and  the  back  is  of  a  rich  metallic  blue;  in  which  respects 
this  species  approaches  the  Java  peacock.  P.  hardwickii  pos- 
sesses a  peculiar  top-knot,  which  is  also  somewhat  like  that  ot 
the  Java  peacock.  In  all  the  species  the  ocelli  on  the  wings 
and  tail  are  either  circular  or  oval,  and  consist  of  a  beautiful, 
iridescent,  greenish-blue  or  greenish-purple  disc,  with  a  black 
border.  This  border  in  P.  chinquis  shades  into  brown,  edged  with 
cream-color,  so  that  the  ocellus  is  here  surrounded  with  variously 
shaded,  though  not  bright,  concentric  zones.  The  unusual  length 
of  the  tail-coverts  is  another  remarkable  character  in  Polyplec- 
tron; for  in  some  of  the  species  they  are  half,  and  in  others  two- 
thirds  as  long  as  the  true  tail-feathers.  The  tail-coverts  are 
ocellated  as  in  the  peacock.  Thus  the  several  species  of  Poly- 
plectron manifestly  make  a  graduated  approach  to  the  peacock 
in  the  length  of  their  tail-coverts,  in  the  zoning  of  the  ocelli, 
and  in  some  other  characters. 

Notwithstanding  this  approach,  the  first  species  of  Polyplectron 
Vv'hich  I  examined  almost  made  me  give  up  the  search;  for  1 
found  not  only  that  the  true  tail-feathers,  which  in  the  peacock 
are  quite  plain,  were  ornamented  with  ocelli,  but  that  the  ocelli 
on  all  the  feathers  differed  fundamentally  from  those  of  the  pea- 
cock, in  there  being  two  on  the  same  feather  (fig.  55),  one  on 
each  side  of  the  shaft.  Hence  I  concluded  that  the  early  pro- 
genitors of  the  peacock  could  not  have  resembled  a  Polyplectron. 
But  on  continuing  my  search,  I  observed  that  in  some  of  the 
species  the  two  ocelli  stood  very  near  each  other;  that  in  the 
tail-feathers  of  P.  hardwickii  they  touched  each  other;  and, 
finally,  that  on  the  tail-coverts  of  this  same  species  as  well  as  of 
P.  malaccense  (fig.  56)  they  were  actually  confluent.  As  the 
central  part  alone  is  confluent,  an  indentation  is  left  at  both  the 
upper  and  lower  ends;  and  the  surrounding  colored  zones  are 
likewise  indented.  A  single  ocellus  is  thus  formed  on  each  tail- 
covert,  though  still  plainly  betraying  its  double  origin.  These 
confluent  ocelli  differ  from  the  single  ocelli  of  the  peacock  in 
having  an  indentation  at  both  ends,  instead  of  only  at  the  lower 
or  basal  end.  The  explanation,  however,  of  this  difference  is 
not  difficult;  in  some  species  of  Polyplectron  the  two  oval  ocelli 
on  the  same  feather  stand  parallel  to  each  other;  in  other  spe- 
cies (as  in  P.  chinquis)  they  converge  towards  one  end;  now  the 
partial  confluence  of  two  convergent  ocelli  would  manifestly  leave 
a  much  deeper  indentation  at  the  divergent  than  at  the  con- 
vergent end.  It  is  also  manifest  that  if  the  convergence  were 
strongly  pronounced  and  the  confluence  complete,  the  indentation 
at  the  convergent  end  would  tend  to  disappear. 


BIRDS— GRADATION    OF    CHARACTERS. 


427 


The  tail-feathers  in  both  species  of  peacock  are  entirely  destitute 
of  ocelli,  and  this  apparently  is  related  to  their  being  covered  up 
and  concealed  by  the  long  tail-coverts.  In  this  respect  they  dif- 
fer remarkably  from  the  tail-feathers  of  Polyplectron,  which  in 
most  of  the  species  are  ornamented  with  larger  ocelli  than  those 
on  the  tail-coverts.  Hence  I  was  led  carefully  to  examine  the 
tail-feathers  of  the  several  species,  in  order  to  discover  whether 


Fig-.  55.  Part  of  a  tail-covert 
of  Polyplectron  chinquis, 
with  the  two  ocelli  of  nat. 

size. 


Fig-.  56.  Part  of  a  tail-covert 
of  Polyplectron  malac- 
cense,  with  the  two  ocelli, 
partially  confluent,  of  nat, 
size. 


their  ocelli  showed  any  tendency  to  disappear;  and  to  my  great 
satisfaction,  this  appeared  to  be  so.  The  central  taii-feathers 
of  P.  napoleonis  have  the  two  ocelli  on  each  side  of  the  shaft  per- 
fectly developed;  but  the  inner  ocellus  becomes  less  and  less 
conspicuous  on  the  more  exterior  tail-feathers,  until  a  mere 
shadow  or  rudiment  is  left  on  the  inner  side  of  the  outermost 
feather.  Again,  in  P.  malaccense,  the  ocelli  on  the  tail-coverts 
are,  as  we  have  seen,  confluent;  and  these  feathers  are  of  un- 
usual length,  being  two-thirds  of  the  length  of  the  tail-feathers. 
so  that  in  both  these  respects  they  approach  the  tail-coverts  of  the 
peacock.  Now  in  P.  malaccense  the  two  central  tail-feathers 
alcne  are  ornamented,  each  with  two  brightly-colored  ocelli,  the 
inner  ocellus  having  completely  disappeared  from  all  the  other 
tail-feathers.  Consequently  the  tail-coverts  and  tail-feathers  of 
this  species  of  Polyplectron  make  a  near  approach  in  structure 
and  ornamentation  to  the  corresponding  feathers  of  the  peacock. 


428  THE  DESCENT  OF  MAN. 

As  far,  then,  as  gradation  throws  light  on  the  steps  by  which 
the  magnificent  train  of  the  peacock  has  been  acquired,  hardly 
anything  more  is  needed.  If  we  picture  to  ourselves  a  progeni- 
tor of  the  peacock  in  an  almost  exactly  intermediate  condition 
between  the  existing  peacock,  with  his  enormously  elongated  tail- 
coverts,  ornamented  with  single  ocelli,  and  an  ordinary  gallina- 
ceous bird  with  short  tail-coverts,  merely  spotted  with  some  color, 
we  shall  see  a  bird  allied  to  Polyplectron — that  is,  with  tail- 
coverts,  capable  of  erection  and  expansion,  ornamented  with  two 
partially  confluent  ocelli,  and  long  enough  almost  to  conceal  the 
tail-feathers,  the  latter  having  already  partially  lost  their  ocelli. 
The  indentation  of  the  central  disc  and  of  the  surroundihg  zones 
of  the  ocellus,  in  both  species  of  peacock,  speaks  plainly  in  favor 
of  this  view,  and  is  otherwise  inexplicable.  The  males  of  Poly- 
plectron are  no  doubt  beautiful  birds,  but  their  beauty,  when 
viewed  from  a  little  distanv-3e,  cannot  be  compared  with  that  of 
the  peacock.  Many  female  progenitors  of  the  peacock  must,  dur- 
ing a  long  line  of  descent,  have  appreciated  this  superiority;  for 
they  have  unconsciously,  by  the  continued  preference  of  the  most 
beautiful  males,  rendered  the  peacock  the  most  splendid  of  living 
birds. 

Argus  Pheasant. — Another  excellent  case  for  investigation  is 
offered  by  the  ocelli  on  the  wing-feathers  of  the  Argus  pheasant, 
which  are  shaded  in  so  wonderful  a  manner  as  to  resemble  balls 
lying  loose  within  sockets,  and  consequently  differ  from  ordinary 
ocelli.  No  one,  I  presume,  will  attribute  the  shading,  which  has 
excited  the  admiration  of  many  experienced  artists,  to  chance 
— to  the  fortuitous  concourse  of  atoms  of  coloring  matter.  That 
these  ornaments  should  have  been  formed  through  the  selection 
of  many  successive  variations,  not  one  of  which  was  originally 
intended  to  produce  the  ball-and-socket  effect,  seems  as  incredible, 
as  that  one  of  Raphael's  Madonnas  should  have  been  formed  by 
the  selection  of  chance  daubs  of  paint  made  by  a  long  succession 
of  young  artists,  not  one  of  whom  intended  at  first  to  draw  the 
human  figure.  In  order  to  discover  how  the  ocelli  have  been  de- 
veloped, we  cannot  look  to  a  long  line  of  progenitors,  nor  to 
many  closely-allied  forms,  for  such  do  not  now  exist.  But  fortu- 
nately the  several  feathers  on  the  wing  suffice  to  give  us  a  clue 
to  the  problem,  and  they  prove  to  demonstration  that  a  gradation 
is  at  least  possible  from  a  mere  spot  to  a  finished  ball-and-socket 
ocellus. 

The  wing-feathers,  bearing  the  ocelli,  are  covered  with  dark 
stripes  (fig.  57)  or  with  rows  of  dark  spots  (fig.  59),  each  stripe 
or  row  of  spots  running  obliquely  down  the  outer  side  of  the 
shaft  to  one  of  the  ocelli.  The  spots  are  generally  elongated  in  a 
line  transverse   to  the  row  in  which   they   stand.    They  often 


BIRDS— GRADATION    OF    CHARACTERS. 


429 


become  confluent,  either  in  the  line  of  the  row— and  then  they 
form  a  longitudinal  stripe— or  transversely,  that  is,  with  the  spots 
in  the  adjoining  rows,  and  then  they  form  transverse  stripes.  A 
spot  sometimes  breaks  up  into  smaller  spots,  which  still  stand  in 
their  proper  places. 

It  will  be  convenient  first 
to  describe  a  perfect  ball- 
and-socket  ocellus.  This 
consists  of  an  intensely 
black  circular  ring,  sur- 
rounding a  space  shaded  so 
as  exactly  to  resemble  a 
ball.  The  figure  here  given 
has  been  admirably  drawn 
by  Mr.  Ford  and  well  en- 
graved, but  a  woodcut  can- 
not exhibit  the  exquisite 
shading  of  the  original. 
The  ring  is  almost  always 
slightly  broken  or  inter- 
rupted (see  fig.  57)  at  a 
point  in  the  upper  half,  a 
little  to  the  right  of,  and 
above  the  white  shade  on 
the  enclosed  ball;  it  is  also 
sometimes  broken  towards 
the  base  on  the  right  hand. 
These  little  breaks  have  an 
important  meaning.  The 
ring  is  always  much  thick- 
ened, with  the  edges  ill-de- 
fined towards  the  left-hand 
upper  corner,  the  feather 
being  held  erect,  in  the  po- 
sition in  which  it  is  here 
drawn.  Beneath  this  thick- 
ened part  there  is  on  the 
surface  of  the  ball  an 
oblique  almost  pure-white 
mark,  which  shades  off 
downwards  into  a  pale- 
leaden  hue,   and  this  into 

yellowish  and  brown  tints,  which  insensibly  become  darker 
and  darker  towards  the  lower  part  of  the  ball.  It  is  this 
shading  which  gives  so  admirably  the  effect  of  light  shining  on 
a  convex  surface.  If  one  of  the  balls  be  examined,  it  will  be  seen 
that  the  lower  part  is  of  a  brown  tint  and  is  indistinctly  sepa- 


Fig-.  57.  Part  of  secondary  wing-feath- 
er of  Arg-us  pheasant,  showing  two 
perfect  ocelli,  a  and  b.  A,  B,  C,  D, 
&c.,  are  dark  stripes  running-  ob- 
liquely down,  each  to  an  ocellus. 

[Much  of  the  web  on  both  sides,  es- 
specially  to  the  left  of  the  shaft,  has 
been  cut  off.] 


430 


THE  DESCENT  OF  MAN. 


rated  by  a  curved  oblique  line  from  the  upper  part,  which  is 
yellower  and  more  leaden;  this  curved  oblique  line  runs  at  right 
angles  to  the  longer  axis  of  the  white  patch  of  light,  and  indeed 
of  all  the  shading;  but  this  difference  in  color,  which  cannot 
of  course  be  shown  in  the  woodcut,  does  not  in  the  least  interfere 
with  the  perfect  shading  of  the  ball.  It  should  be  particularly 
observed  that  each  ocellus  stands  in  obvious  connection  either 
with  a  dark  stripe,  or  with  a  longitudinal  row  of  dark  spots,  for 
both  occur  indifferently  on  the  same  feather.  Thus  in  fig.  57  stripe 
A  runs  to  ocellus  a;  B  runs  to  ocellus  b;  stripe  C  is  broken  in 
the  upper  part,  and  runs  down  to  the  next  succeeding  ocellus,  not 
represented  in  the  woodcut;  D  to  the  next  lower  one,  and  so  with 
the  stripes  E  and  F.    Lastly,  the  ocelli  are  separated  from  each 

other  by  a  pale  surface  bear- 
ing irregular  black  marks. 

I  will  next  describe  the 
other  extreme  of  the  series, 
namely,  the  first  trace  of  an 
ocellus.  The  short  secondary 
wing- feather  (fig.  58),  nearest 
to  the  body,  is  marked  like 
the  other  feathers,  with 
oblique,  longitudinal,  rather 
irregular,  rows  of  very  dark 
spots.  The  basal  spot,  or  that 
nearest  the  shaft,  in  the  five 
lower  rows  (excluding  the 
lowest  one)  is  a  little  larger 
than  the  other  spots  of  the 
same  row,  and  a  little  more 
elongated  in  a  transverse  di- 
rection. It  differs  also  from 
the  other  spots  by  being  bor- 
dered on  its  upper  side  with 
some  dull  fulvous  shading. 
But  this  spot  is  not  in  any 
way  more  remarkable  than 
those  on  the  plumage  of 
many  birds,  and  might  easily 
be  overlooked.  The  next  higher  spot  does  not  differ  at  all  from 
the  upper  ones  in  the  same  row.  The  larger  basal  spots  occupy 
exactly  the  same  relative  position  on  these  feathers,  as  do  the 
perfect  ocelli  on  the  longer  wing-feathers. 

By  looking  to  the  next  two  or  three  succeeding  wing-feathers, 
an  absolutely  insensible  gradation  can  be  traced  from  one  of  the 
last-described  basal  spots,  together  with  the  next  higher  one  in 
the  same  row,  to  a  curious  ornament,  which  cannot  be  called  an 


Fig.  58.  Basal  part  of  the  sec- 
ondary-wing' feather,  nearest 
to  the  body. 


BIRDS— GRADATION    OF    CHARACTERS. 


431 


ocellus,  and  which  I  will  name,  from  the  want  of  a  better  term, 
an  "elliptic  ornament."  These  are  shown  in  the  accompanying 
figure  (ng.  59).  We  here  see  several  oblique  rows,  A,  B,  C,  D, 
&c.  (see  the  lettered  diagram  on  the  right  hand),  of  dark  spots 
of  the  usual  character.  Each  row  of  spots  runs  down  to  and  is 
connected  with  one  of  the  elliptic  ornaments,  in  exactly  the 
same  manner  as  each  stripe  in  fig.  57  runs  down  to,  and  is  con- 
nected with  one  of  the  ball-and-socket  ocelli.    Looking  to  any 


Pig.  59.  Portion  of  one  of  the  secondary  wing--feathers  near  to  the 
body,  showing  the  so-called  elliptic  ornaments.  The  right-hand  fig- 
ure is  given  merely  as  a  diagram  for  the  sake  of  the  letters  of 
reference. 

A,  B,  C,  D,  &c.  Rows  of  spots  running  down  to  and  forming  the  ellip- 
tic ornaments. 

b.  Lowest  spot  or  mark  in  row  B. 

c.  The  next  succeeding  spot  or  mark  in  the  same  row. 

d.  Apparently  a  broken  prolongation  of  the  spot  c  in  the  same  row  B. 

one  row,  for  instance,  B,  in  fig.  59,  the  lowest  mark  (b)  is  thicker 
and  considerably  longer  than  the  upper  spots,  and  has  its  left 
extremity  pointed  and  curved  upwards.  This  black  mark  is 
abruptly  bordered  on  its  upper  side  by  a  rather  broad  space  of 
richly  shaded  tints,  beginning  with  a  narrow  brown  zone,  which 
passes  into  orange,  and  this  into  a  pale  leaden  tint,  with  the  end 
towards  the  shaft  much  paler.    These  shaded  tints  together  fill  up 


432  THE  DESCENT  OF  MAN. 

the  whole  inner  space  of  the  elliptic  ornament.  The  mark  (b) 
corresponds  in  every  respect  with  the  basal  shaded  spot  of  the 
simple  feather  described  in  the  last  paragraph  (fig.  58),  but  is 
more  highly  developed  and  more  brightly  colored.  Above  and  to 
the  right  of  this  spot  (b  fig.  59),  with  its  bright  shading,  there  is 
a  long  narrow,  black  mark  (c),  belonging  to  the  same  row,  and 
which  is  arched  a  little  downwards  so  as  to  face  (b).  This  mark 
is  sometimes  broken  into  two  portions.  It  is  also  narrowly  edged 
on  the  lower  side  with  a  fulvous  tint.  To  the  left  of  and  above 
c,  in  the  same  oblique  direction,  but  always  more  or  less  distinct 
from  it,  there  is  another  black  mark  (d).  This  mark  is  generally 
sub-triangular  and  irregular  in  shape,  but  in  the  one  lettered  in 
the  diagram  it  is  unusually  narrow,  elongated,  and  regular.  It 
apparently  consists  of  a  lateral  and  broken  prolongation  of  the 
mark  (c),  together  with  its  confluence  with  a  broken  and  pro- 
longed part  of  the  next  spot  above;  but  I  do  not  feel  sure  of  this. 
These  three  marks,  b,  c,  and  d,  with  the  intervening  bright  shades, 
form  together  the  so-called  elliptic  ornament.  These  ornaments 
placed  parallel  to  the  shaft,  manifestly  correspond  in  position 
with  the  ball-and-socket  ocelli.  Their  extremely  elegant  ap- 
pearance cannot  be  appreciated  in  the  drawing,  as  the  orange  and 
leaden  tints,  contrasting  so  well  with  the  black  marks,  cannot  be 
shown. 

Between  one  of  the  elliptic  ornaments  and  a  perfect  ball-and- 
socket  ocellus,  the  gradation  is  so  perfect  that  it  is  scarcely  pos- 
sible to  decide  when  the  latter  term  ought  to  be  used.  The  pas- 
sage from  the  one  into  the  other  is  effected  by  the  elongation, 
and  greater  curvature  in  opposite  directions  of  the  lower  black 
mark  (b  fig.  59),  and  more  especially  of  the  upper  one  (c),  to- 
gether with  the  contraction  of  the  elongated  sub-triangular  or 
narrow  mark  (d),  so  that  at  last  these  three  marks  become  con- 
fluent, forming  an  irregular  elliptic  ring.  This  ring  is  gradually 
rendered  more  and  more  circular  and  regular,  increasing  at  the 
same  time  in  diameter.  I  have  here  given  a  drawing  (fig.  60) 
of  the  natural  size  of  an  ocellus  not  as  yet  quite  perfect.  The 
lower  part  of  the  black  ring  is  much  more  curved  than  is  the  lower 
mark  in  the  elliptic  ornament  (b  fig.  59).  The  upper  part  of 
the  ring  consists  of  two  or  three  separate  portions;  and  there  is 
only  a  trace  of  the  thickening  of  the  portion  which  forms  the 
black  mark  above  the  white  shade.  This  white  shade  itself  is 
not  as  yet  much  concentrated;  and  beneath  it  the  surface  is 
brighter  colored  than  ^n  a  perfect  ball-and-socket  ocellus.  Even 
in  the  most  perfect  ocelli,  traces  of  the  junction  of  three  or  four 
elongated  black  marks,  by  which  the  ring  has  been  formed,  may 
often  be  detected.    The  irregular  sub-triangular  or  narrow  mark 


BIRDS— GRADATION    OF    CHARACTERS. 


433 


(d.  fig.  59),  manifestly  forms,  by  its  contraction  and  equalization, 
the  thickened  portion  of  the  ring  above  the  white  shade  on  a  per- 
fect ball-and-socket  ocellus.  The  lower  part  of  the  ring  is  invari- 
ably a  little  thicker  than  the  other 
parts  (see  fig.  57),  and  this  follows 
from  the  lower  black  mark  of  the 
elliptic  ornament  (b  fig.  59)  having 
originally  been  thicker  than  the 
upper  mark  (c).  Every  step  can  be 
followed  in  the  process  of  conflu- 
ence and  modification;  and  the 
black  ring  which  surrounds  the  ball 
of  the  ocellus  is  unquestionably 
formed  by  the  union  and  modifica- 
tion of  the  three  black  marks,  b,  c, 
d,  of  the  elliptic  ornament.  The 
irregular  zigzag  black  marks  be- 
tween the  successive  ocelli  (see 
again  fig.  57)  are  plainly  due  to  the 
breaking  up  of  the  somewhat  mora 
regular  but  similar  marks  between 
the  elliptic  ornaments. 

The  successive  steps  in  the  shad- 
ing of  the  ball-and-socket  ocelli  can 
be  followed  out  with  equal  clear- 
ness. The  brown,  orange,  and 
pale  leaden  narrow  zones,  which 
border  the  lower  black  mark  of 
the  elliptic  ornament,  can  be  seen  gradually  to  become  more 
and  more  softened  and  shaded  into  each  other,  with  the 
upper  lighter  part  towards  the  left-hand  corner  rendered  still 
lighter,  so  as  to  become  almost  white,  and  at  the  same  time  more 
contracted.  But  even  in  the  most  perfect  ball-and-socket  ocelli 
a  slight  difference  in  the  tints,  though  not  in  the  shading,  be- 
tween the  upper  and  lower  parts  of  the  ball  can  be  perceived,  as 
before  noticed;  and  the  line  of  separation  is  oblique,  in  the 
same  direction  as  the  bright-colored  shades  of  the  elliptic  orna- 
ments. Thus  almost  every  minute  detail  in  the  shape  and  color- 
ing of  the  ball-and-socket  ocelli  can  be  shown  to  follow  from 
gradual  changes  in  the  elliptic  ornaments;  and  the  development 
of  the  latter  can  be  traced  by  equally  small  steps  from  the  union 
of  two  almost  simple  spots,  the  lov/er  one  (fig.  58)  having  some 
dull  fulvous  shading  on  its  upper  side. 

The  extremities  of  the  longer  secondary  feathers  which  bear 
the  perfect  ball-and-socket  ocelli,  are  peculiarly  ornamented  (fig. 
61).     The  oblique  longitudinal  stripes  suddenly  cease  upwards 

29 


Fig-.  60.  An  ocellus  in  an 
intermediate  condition  be- 
tween the  elliptic  orna- 
ment and  the  perfect  ball- 
and-socket  ocellus. 


434 


THE  DESCENT  OF   MAN. 


and  become  confused;  and  above  this  limit  the  whole  upper  end 
of  the  feather  (a)  is  covered  with  white  dots,  surrounded  by 
little  black  rings,  standing  on  a  dark  ground.     The  oblique  stripe 

belonging  to  the  uppermost  ocellus 
(b)  is  barely  represented  by  a  very 
short  irregular  black  mark  with  the 
usual,  curved,  transverse  base.  As 
this  stripe  is  thus  abruptly  cut  off, 
we  can  perhaps  understand  from 
what  has  gone  before,  how  it  is  that 
the  upper  thickened  part  of  the  ring 
is  here  absent;  for,  as  before 
stated,  this  thickened  part  appar- 
ently stands  in  some  relation  with 
a  broken  prolongation  from  the  next 
higher  spot.  From  the  absence  of 
the  upper  and  thickened  part  of  the 
ring,  the  uppermost  ocellus,  though 
perfect  in  all  other  respects,  ap- 
pears as  if  its  top  had  been  obliquely 
sliced  off.  It  would,  I  think,  per- 
plex any  one,  who  believes  that  the 
plumage  of  the  Argus-pheasant  was 
created  as  we  now  see  it,  to  account 
for  the  imperfect  condition  of  the 
uppermost  ocellus.  I  should  add 
that  on  the  secondary  wing-feather 
farthest  from  the  body,  all  the 
ocelli  are  smaller  and  less  perfect 
than  on  the  other  feathers,  and 
have  the  upper  part  of  the  ring  de- 
ficient, as  in  the  case  just  men- 
Portion  near  summit  tioned.  The  imperfection  here 
seems  to  be  connected  with  the  fact 
that  the  spots  on  this  feather  show 
less  tendency  than  usual  to  become 
confluent  into  stripes;  they  are  on 
the  contrary,  often  broken  up  into 
shading-  above  the  white  smaller  spots,  so  that  two  or  three 
mark  on  the  summit  of  rov/s  run  dowu  to  the  same 
the  ocellus  is  here  a  little    ocellus. 

c  Perfect^ocellus  There  still  remains  another  very 

curious    point,    first    observed    by 
Mr.  T.  W.  Woods,^^  which  deserves  attention.    In  a  photograph. 


Fig-.  6! 

of  one  of  the  secondary 
wing-feathers,  bearing  per- 
fect ball-and-socket  ocelli. 

a.  Ornamented  upper  part. 

b.  Uppermost,   imperfect  ball- 
and-socket  ocellus.    (The 


51  The  'Field,'  May  28,  1870. 


BIRDS— GRADATION    OF    CHARACTERS.  435 

given  me  by  Mr.  Ward,  of  a  specimen  mounted  as  in  the  act  of 
display,  it  may  be  seen  that  on  the  feathers  which  are  held  per- 
pendicularly, the  white  marks  on  the  ocelli,  representing  light 
reflected  from  a  convex  surface^  are  at  the  upper  or  further  end, 
that  is,  are  directed  upwards;  and  the  bird  whilst  displaying 
himself  on  the  ground  would  naturally  be  illuminated  from  above. 
But  here  comes  the  curious  point,  the  outer  feathers  are  held 
almost  horizontally,  and  their  ocelli  ought  likewise  to  appear  as 
if  illuminated  from  above,  and  consequently  the  white  marks  ought 
to  be  placed  on  the  upper  sides  of  the  ocelli;  and  wonderful  as  is 
the  fact  they  are  thus  placed!  Hence  the  ocelli  on  the  several 
feathers,  though  occupying  very  different  positions  with  respect 
to  the  light,  all  appear  as  if  illuminate^  from  above,  just  as  an 
artist  would  have  shaded  them.  Nevertheless  they  are  not  il- 
luminated from  strictly  the  same  point  as  they  ought  to  be;  for 
the  white  marks  on  the  ocelli  of  the  feathers  which  are  held 
almost  horizontally,  are  placed  rather  too  much  towards  the 
further  end;  that  is  they  are  not  sufficiently  lateral.  We  have, 
however,  no  right  to  expect  absolute  perfection  in  a  part  rendered 
ornamental  through  sexual  selection,  any  more  than  we  have 
in  a  part  modified  through  natural  selection  for  real  use;  for 
instance  in  that  wondrous  organ  the  human  eye.  And  we  know 
what  Helmholtz,  the  highest  authority  in  Europe  on  the  subject, 
has  said  about  the  human  eye;  that  if  an  optician  had  sold  him 
an  instrument  so  carelessly  made,  he  would  have  thought  himself 
fully  justified  in  returning  it.^- 

We  have  now  seen  that  a  perfect  series  can  be  followed,  from 
simple  spots  to  the  wonderful  ball-and-socket  ornaments.  Mr. 
Gould,  who  kindly  gave  me  some  of  these  feathers,  fully  agrees 
with  me  in  the  completeness  of  the  gradation.  It  is  obvious 
that  the  stages  in  development  exhibited  by  the  feathers  on  the 
same  bird,  do  not  at  all  necessarily  show  us  the  steps  passed 
through  by  the  extinct  progenitors  of  the  species;  but  they  prob- 
ably give  us  the  clue  to  the  actual  steps,  and  they  at  least  prove 
to  demonstration  that  a  gradation  is  possible.  Bearing  in  mind 
how  carefully  the  male  Argus  pheasant  displays  his  plumes  before 
the  female,  as  well  as  the  many  facts  rendering  it  probable  that 
female  birds  prefer  the  more  attractive  males,  no  one  who  ad- 
mits the  agency  of  sexual  selection  in  any  case,  will  deny  that 
a  simple  dark  spot  with  some  fulvous  shading  might  be  converted, 
through  the  approximation  and  modification  of  two  adjoining 
spots,  together  with  some  slight  increase  of  color,  into  one  of  the 
so-called  elliptic  ornaments.     These  latter  ornaments  have  been 

52  'Popular  Lectures  on  Scientific  Subjects,'  Eng.  trans.  1873,  pp.  219, 
227,  269,  390. 


435  THE  DESCENT  OF  MAN. 

shown  to  many  persons,  and  all  have  admitted  that  they  are 
beautiful,  some  thinking  them  even  more  so  than  the  ball-and- 
socket  ocelli.  As  the  secondary  plumes  become  lengthened 
through  sexual  selection,  and  as  the  elliptic  ornaments  increased 
in  diameter,  their  colors  apparently  became  less  bright;  and 
then  the  ornamentation  of  the  plumes  had  to  be  gained  by  an 
improvement  in  the  pattern  and  shading;  and  this  process  was 
carried  on  until  the  wonderful  ball-and-socket  ocelli  were  finally 
developed.  Thus  we  can  understand — and  in  no  other  way  as 
it  seems  to  me — the  present  condition  and  origin  of  the  orna- 
ments on  the  wing-feathers  of  the  Argus  pheasant. 

From  the  light  afforded  by  the  principle  of  gradation — from 
what  we  know  of  the  laws  of  variation — from  the  changes  which 
have  taken  place  in  many  of  our  domesticated  birds — and,  lastly, 
from  the  character  (as  we  shall  hereafter  see  more  clearly)  of  the 
immature  plumage  of  young  birds — we  can  sometimes  indicate 
with  a  certain  amount  of  confidence,  the  probable  steps  by  which 
the  males  have  acquired  their  brilliant  plumage  and  various  or- 
naments; yet  in  many  cases  we  are  involved  in  complete  darkness. 
Mr.  Gould  several  years  ago  pointed  out  to  me  a  humming-bird, 
the  Urosticte  benjamini,  remarkable  for  the  curious  differences 
between  the  sexes.  The  male,  besides  a  splendid  gorget,  has 
greenish-black  tail-feathers,  with  the  four  central  ones  tipped 
with  white;  in  the  female,  as  with  most  of  the  allied  species, 
the  three  outer  tail-feathers  on  each  side  are  tipped  with  white, 
so  that  the  male  has  the  four  central,  whilst  the  female  has  the 
six  exterior  feathers  ornamented  with  white  tips.  What  makes 
the  case  more  curious  is  that,  although  the  coloring  of  the  tail 
differs  remarkably  in  both  sexes  of  many  kinds  of  humming- 
birds, Mr.  Gould  does  not  know  a  single  species,  besides  the 
Urosticte,  in  which  the  male  has  the  four  central  feathers  tipped 
with  white. 

The  Duke  of  Argyll,  in  commenting  on  this  case,^^  passes  over 
sexual  selection,  and  asks,  "What  explanation  does  the  law  of 
"natural  selection  give  of  such  specific  varieties  as  these?"  He 
answers  "none  whatever;"  and  I  quite  agree  with  him.  But 
can  this  be  so  confidently  said  of  sexual  selection?  Seeing  in 
how  many  ways  the  tail-feathers  of  bumming-birds  differ,  why 
should  not  the  four  central  feathers  have  varied  in  this  one  spe» 
cies  alone,  so  as  to  have  acquired  white  tips?  The  variations 
may  have  been  gradual,  or  somewhat  abrupt  as  in  the  case  re- 
cently given  of  the  humming-birds  near  Bogota,  in  wnich  certain 
individuals  alone  have  the  "central  tail-feathers  tipped  with  beau- 

83  'The  Reign  of  Law,'  1867,  p.  247. 


BIRDS— GRADATION    OF    CHARACTERS.  437 

"tifiil  green."  In  the  female  of  the  Urosticte  I  noticed  extremely 
minute  or  rudimental  white  tips  to  the  two  outer  of  the  four  cen- 
tral black  tail-feathers,  so  that  here  we  have  an  indication  of 
change  of  some  kind  in  the  plumage  of  this  species.  If  we  grant  the 
possibility  of  the  central  tail-feathers  of  the  male  varying  in 
whiteness,  there  is  nothing  strange  in  such  variations  having 
been  sexually  selected.  The  white  tips,  together  vs^ith  the  small 
white  ear-tufts,  certainly  add,  as  the  Duke  of  Argyll  admits,  to 
the  beauty  of  the  male;  and  whiteness  is  apparently  appreciated 
by  other  birds,  as  may  be  inferred  from  such  cases  as  the  snow- 
white  male  of  the  Bell-bird.  The  statement  made  by  Sir  R. 
Heron  should  not  be  forgotten,  namely,  that  his  peahens,  when 
debarred  from  access  to  the  pied  peacock,  would  not  unite  with 
any  other  male,  and  during  that  season  produced  no  offspring. 
Nor  is  it  strange  that  variations  in  the  tail-feathers  of  the  Uros- 
ticte should  have  been  specially  selected  for  the  sake  of  ornament, 
for  the  next  succeeding  genus  in  the  family  takes  its  name  of 
Metallura  from  the  splendor  of  these  feathers.  We  have,  more- 
over, good  evidence  that  humming-birds  take  especial  pains  in 
displaying  their  tail-feathers;  Mr.  Belt,^*  after  describing  the 
beauty  of  the  Florisuga  mellivora,  says,  "I  have  seen  the  female 
"sitting  on  a  branch,  and  two  males  displaying  their  charms  in 
"front  of  her.  One  would  shoot  up  like  a  rocket,  then  suddenly 
"expanding  the  snow-white  tail,  like  an  inverted  parachute, 
"slowly  descend  in  front  of  her,  turning  round  gradually  to  show 

"off  back  and  front The  expanded  white  tail  covered 

"more  space  than  all  the  rest  of  the  bird,  and  was  evidently  the 
"grand  feature  in  the  performance.  Whilst  one  male  was  de- 
"scending,  the  other  would  shoot  up  and  come  slowly  down  ex- 
"panded.  The  entertainment  would  end  in  a  fight  between  the 
"two  performers;  but  whether  the  most  beautiful  or  the  most  pug- 
"nacious  was  the  accepted  suitor,  I  know  not."  Mr.  Gould,  after 
describing  the  peculiar  plumage  of  the  Urosticte,  adds,  "that  orna- 
"ment  and  variety  is  the  sole  object,  I  have  myself  but  little 
"doubt. "°^  If  this  be  admitted,  we  can  perceive  that  the  males 
which  during  former  times  were  decked  in  the  most  elegant  and 
novel  manner  would  have  gained  an  advantage,  not  in  the  or- 
dinary struggle  for  life,  but  in  rivalry  with  other  males,  and 
would  have  left  a  larger  number  of  offspring  to  inherit  their  new- 
ly-acquired beauty. 


5^  'The  Naturalist  in  Nicaragua,'  1874,  p.  112. 
65  'Introduction  to  the  Trochilidae,'  1861,  p.  110. 


29 


438  THE  DESCENT  OF  MAN. 


CHAPTER    XV. 

BIRDS— Continued. 

Discussion  as  to  why  the  males  alone  of  some  species,  and  both  sexes 
of  others,  are  brightly  colored— On  sexually-limited  inheritance,  as 
applied  to  various  structures  and  to  brightly-colored  plumage— 
Nidification  in  relation  to  color— Loss  of  nuptial  plumage  during 
the  winter. 

We  have  in  this  chapter  to  consider,  why  the  females  of  many 
birds  have  not  acquired  the  same  ornaments  as  the  male;  and 
why,  on  he  other  hand,  both  sexes  of  many  other  birds  are 
equally,  or  almost  equally,  ornamented?  In  the  following  chap- 
ter we  shall  consider  the  few  cases  in  which  the  female  is  more 
conspicuously  colored  than  the  male. 

In  my  'Origin  of  Species'^  I  briefly  suggested  that  the  long  tail 
of  the  peacock  would  be  inconvenient,  and  the  conspicuous  black 
color  of  the  male  capercailzie  dangerous,  to  the  female  during 
the  period  of  incubation;  and  consequently  that  the  transmission 
of  these  characters  from  the  male  to  the  female  offspring  had 
been  checked  through  natural  selection.  I  still  think  that  this 
may  have  occurred  in  some  few  instances:  but  after  mature  re- 
flection on  all  the  facts  which  I  have  been  able  to  collect,  I  am 
now  inclined  to  believe  that  when  the  sexes  differ,  the  successive 
variations  have  generally  been  from  the  first  limited  in  their 
transmission  to  the  same  sex  in  which  they  first  arose.  Since 
my  remarks  appeared,  the  subject  of  sexual  coloration  has  been 
discussed  in  some  very  interesting  papers  by  Mr.  Wallace,-  who 
believes  that  in  almost  all  cases  the  successive  variations  tended 
at  first  to  be  transmitted  equally  to  both  sexes;  but  that  the 
female  was  saved,  through  natural  selection,  from  acquiring  the 
conspicuous  colors  of  the  male,  owing  to  the  danger  which  she 
would  thus  have  incurred  during  incubation. 

This  view  necessitates  a  tedious  discussion  on  a  difficult  point. 


1  Fourth  edition,  1866,  p.  241. 

2  'Westminster  Review,'  July,  1867.    'Journal  of  Travel,'  vol.  i.  1868, 
..  73. 


BIRDS-SEXUALLY  LIMITED  INHERITANCE.  439 

namely,  whettier  the  transmission  of  a  character,  which  is  at 
first  inherited  by  both  sexes,  can  be  subsequently  limited  in  its 
transmission  to  one  sex  alone  by  means  of  natural  selection.  We 
must  bear  in  mind,  as  shown  in  the  preliminary  chapter  on  sexual 
selection,  that  characters  which  are  limited  in  their  development 
to  one  sex  are  always  latent  in  the  other.  An  imaginary  illustra- 
tion will  best  aid  us  in  seeing  the  difficulty  of  the  case:  we  may 
suppose  that  a  fancier  wished  to  make  a  breed  of  pigeons,  in 
which  the  males  alone  should  be  colored  of  a  pale  blue,  whilst  the 
females  retained  their  former  slaty  tint.  As  with  pigeons  char- 
acters of  all  kinds  are  usually  transmitted  to  both  sexes  equally, 
the  fancier  would  have  to  try  to  convert  this  latter  form  of  in- 
heritance into  sexually-limited  transmission.  All  that  he  could 
do  would  be  to  persevere  in  selecting  every  male  pigeon  which 
was  in  the  least  degree  of  a  paler  blue;  and  the  natural  result 
of  this  process,  if  steadily  carried  on  for  a  long  time,  and  if  the 
pale  variations  were  strongly  inherited  or  often  recurred,  would 
be  to  make  his  whole  stock  of  a  lighter  blue.  But  our  fancier 
would  be  compelled  to  match,  generation  after  generation,  his 
pale  blue  males  with  slaty  females,  for  he  wishes  to  keep  the 
latter  of  this  color.  The  result  would  generally  be  the  production 
either  of  a  mongrel  piebald  lot,  or  more  probably  the  speedy  and 
complete  loss  of  the  pale-blue  tint;  for  the  primordial  slaty 
color  would  be  transmitted  with  prepotent  force.  Supposing, 
however,  that  some  pale-blue  males  and  slaty  females  were  pro- 
duced during  each  successive  generation,  and  were  always  crossed 
together;  then  the  slaty  females  would  have,  if  I  may  use  the 
expression,  much  blue  blood  in  their  veins,  for  their  fathers, 
grandfathers,  &c.,  will  all  have  been  blue  birds.  Under  these  cir- 
cumstances it  is  conceivable  (though  I  know  of  no  distinct  facts 
rendering  it  probable)  that  the  slaty  females  might  acquire  so 
strong  a  latent  tendency  to  pale-blueness,  that  they  would  not 
destroy  this  color  in  their  male  offspring,  their  female  offspring 
still  inheriting  the  slaty  tint.  If  so  the  desired  end  of  making  a 
breed  with  the  two  sexes  permanently  different  in  color  might 
be  gained. 

The  extreme  importance,  or  rather  necessity  in  the  above  case 
of  the  desired  character,  namely,  pale-blueness,  being  present 
though  in  a  latent  state  in  the  female,  so  that  the  male  offspring 
should  not  be  deteriorated,  will  be  best  appreciated  as  follows: 
the  male  of  Scemmerring's  pheasant  has  a  tail  thirty-seven 
inches  in  length,  whilst  that  of  the  female  is  only  eight  inches; 
the  tail  of  the  male  common  pheasant  is  about  twenty  inches, 
and  that  of  the  female  twelve  inches  long.  Now  if  the  female 
Soemmerring  pheasant  with  her  short  tail  were  crossed  with  the 
male  common  pheasant,  there  can  be  no  doubt  that  the  male 


'440  THE  DESCENT  OF  MAN. 

hybrid  offspring  would  have  a  mucli  longer  tail  than  that  of  the 
pure  offspring  of  the  common  pheasant.  On  the  other  hand,  it 
the  female  common  pheasant,  with  a  tail  much  longer  than  that 
of  the  female  Scemmerring  pheasant,  were  crossed  with  the  male 
of  the  latter,  the  male  hybrid  offspring  would  have  a  much 
shorter  tail  than  that  of  the  pure  offspring  of  Soemmerring's 
pheasant.^ 

Our  fancier,  in  order  to  make  his  new  breed  with  the  males 
of  a  pale-blue  tint,  and  the  females  unchanged,  would  have  to 
continue  selecting  the  males  during  many  generations;  and  each 
stage  of  paleness  would  have  to  be  fixed  in  the  males,  and  ren- 
dered latent  in  the  females.  The  task  would  be  an  extremely 
difficult  one,  and  has  never  been  tried,  but  might  possibly  be  suc- 
cessfully carried  out.  The  chief  obstacle  would  be  the  early  and 
complete  loss  of  the  pale-blue  tint,  from  the  necessity  of  reiterated 
crosses  with  the  slaty  female,  the  latter  not  having  at  first  any 
latent  tendency  to  produce  pale-blue  offspring. 

On  the  other  hand,  if  one  or  two  males  were  to  vary  ever  so 
slightly  in  paleness,  and  the  variations  were  from  the  first 
limited  in  their  transmission  to  the  male  sex,  the  task  of  making 
a  new  breed  of  the  desired  kind  would  be  easy,  for  such  males 
would  simply  have  to  be  selected  and  matched  with  ordinary 
females.  An  analogous  case  has  actually  occurred,  for  there  are 
breeds  of  the  pigeon  in  Belgium*  in  which  the  males  alone  are 
marked  with  black  striae.  So  again  Mr.  Tegetmeier  has  recently 
shown^  that  dragons  not  rarely  produce  silver-colored  birds, 
which  are  almost  always  hens;  and  he  himself  has  bred  ten 
such  females.  It  is  on  the  other  hand  a  very  unusual  event  when 
a  silver  male  is  produced;  so  that  nothing  would  be  easier,  if 
desired,  than  to  make  a  breed  of  dragons  with  blue  males  and 
silver  females.  This  tendency  is  indeed  so  strong  that  when  Mr. 
'Tegetmeier  at  last  got  a  silver  male  and  matched  him  with  one 
of  the  silver  females,  he  expected  to  get  a  breed  with  both  sexes 
thus  colored;  he  was  however  disappointed,  for  the  young  male 
reverted  to  the  blue  color  of  his  grandfather,  the  young  female 
alone  being  silver.  No  doubt  with  patience  this  tendency  to  re- 
version in  the  males,  reared  from  an  occasional  silver  male 
matched  with  a  silver  hen,  might  be  eliminated,  and  then  both 
sexes  would  be  colored  alike;    and  this  very  process  has  been 

3  Temminck  says  that  the  tail  of  the  female  Phasianus  Soemmer- 
ringii  is  only  six  inches  long,  'Planches  coloriees,'  vol.  v.  1838,  pp.  487 
and  488:  the  measurements  above  given  were  made  for  me  by  Mr. 
Sclater.  For  the  common  pheasant,  see  Macgillivray,  'Hist.  Brit. 
Birds,'  vol.  i.  pp.  118-121. 

*  Dr.  Chapuis,  'Le  Pigeon  Voyageur  Beige,'  1865,  p.  87. 

6  The  'Field,'  Sept.  1872. 


BIRDS-SEXUALLY  LIMITED  INHERITANCE.  441 

followed  witli  success  by  Mr.  Esquilant  in  the  case  of  silver  tur- 
bits. 

With  fowls,  variations  of  color,  limited  in  their  transmission 
to  the  male  sex,  habitually  occur.  When  this  form  of  inheritance 
prevails,  it  might  well  liappen  that  some  of  the  successive  varia- 
tions would  be  transferred  to  the  female,  who  would  then  slightly 
resemble  the  male,  as  actually  occurs  in  some  breeds.  Or  again, 
the  greater  number,  but  not  all,  of  the  successive  steps  might  be 
transferred  to  both  sexes,  and  the  female  would  then  closely  re- 
semble the  male.  There  can  hardly  be  a  doubt  that  this  is  the 
cause  of  the  male  pouter  pigeon  having  a  somewhat  larger  crop, 
and  of  the  male  carrier  pigeon  having  somewhat  larger  wattles, 
than  their  respective  females;  for  fanciers  have  not  selected 
one  sex  more  than  the  other,  and  have  had  no  wish  that  these 
characters  should  be  more  strongly  displayed  in  the  male  than 
in  the  female,  yet  this  is  the  case  with  both  breeds. 

The  same  process  would  have  to  be  followed,  and  the  same 
difficulties  encountered,  if  it  were  desired  to  make  a  breed  with 
the  females  alone  of  some  new  color. 

Lastly,  our  fancier  might  wish  to  make  a  breed  with  the  two 
sexes  differing  from  each  other,  and  both  from  the  parent-species. 
Here  the  difficulty  would  be  extreme,  unless  the  successive  varia- 
tions were  from  the  lirst  sexually  limited  on  both  sides,  and 
then  there  would  be  no  difficulty.  We  see  this  with  the  fowl; 
thus  the  two  sexes  of  the  penciled  Hamburghs  differ  greatly 
from  each  other,  and  from  the  two  sexes  of  the  aboriginal  Gallus 
bankiva;  and  both  are  now  kept  constant  to  their  standard  of 
excellence  by  continued  selection,  which  would  be  impossible  un- 
less the  distinctive  characters  of  both  were  limited  in  their  trans- 
mission. The  Spanish  fowl  offers  a  more  curious  case;  the  male 
has  an  immense  comb,  but  some  of  the  successive  variations,  by 
the  accumulation  of  which  it  was  acquired,  appear  to  have  been 
transferred  to  the  female;  for  she  has  a  comb  many  times  larger 
than  that  of  the  females  of  the  parent-species.  But  the  comb  of 
the  female  differs  in  one  respect  from  that  of  the  male,  for  it  is 
apt  to  lop  over;  and  within  a  recent  period  it  has  been  ordered 
by  the  fancy  that  this  should  always  be  the  case,  and  success  has 
quickly  followed  the  order.  Now  the  lopping  of  the  comb  must 
be  sexually  limited  in  its  transmission,  otherwise  it  would  pre- 
vent the  comb  of  the  male  from  being  perfectly  upright,  which 
would  be  abhorrent  to  every  fancier.  On  the  other  hand,  the  up- 
rightness of  the  comb  in  the  male  must  likewise  be  a  sexually- 
limited  character,  otherwise  it  would  prevent  the  comb  of  the  fe- 
male from  lopping  over. 

From  the  foregoing  illustrations,  we  see  that  even  with  almost 
unlimited  time  at  command,  it  would  be  an  extremely  difficult 


442  THE   DESCENT  OF  MAN. 

and  complex,  perhaps  an  impossible  process,  to  change  one  form 
of  transmission  into  the  other  through  selection.  Therefore, 
without  distinct  evidence  in  each  case,  I  am  unwilling  to  admit 
that  this  has  been  effected  in  natural  species.  On  the  other  hand, 
by  means  of  successive  variations,  which  were  from  the  first 
sexually  limited  in  their  transmission,  there  would  not  be  the 
least  difficulty  in  rendering  a  male  bird  widely  different  in  color 
or  in  any  other  character  from  the  female;  the  latter  being  left 
unaltered,  or  slightly  altered,  or  specially  modified  for  the  sake 
of  protection. 

As  bright  colors  are  of  service  to  the  males  in  their  rivalry 
with  other  males,  such  colors  would  be  selected,  whether  or  not 
they  were  transmitted  exclusively  to  the  same  sex.  Consequently 
the  females  might  be  expected  often  to  partake  of  the  brightness 
of  the  males  to  a  greater  or  less  degree;  and  this  occurs  with 
a  host  of  species.  If  all  the  successive  variations  were  transmitted 
equally  to  both  sexes,  the  females  would  be  indistinguishable 
from  the  males;  and  this  likewise  occurs  with  many  birds.  If, 
however,  dull  colors  were  of  high  importance  for  the  safety  of 
the  female  during  incubation,  as  with  many  ground  birds,  the 
females  which  varied  m  brightness,  or  which  received  through 
inheritance  from  the  males  any  marked  accession  of  brightness, 
would  sooner  or  later  be  destroyed.  But  the  tendency  in  the 
males  to  continue  for  an  indefinite  period  transmitting  to  their 
female  offspring  their  own  brightness,  would  have  to  be  elim- 
inated by  a  change  in  the  form  of  inheritance;  and  this,  as  shown 
by  our  previous  illustration,  would  be  extremely  difficult.  The 
more  probable  result  of  the  long-continued  destruction  of  the 
more  brightly-colored  females,  supposing  the  equal  form  of  trans- 
mission to  prevail,  would  be  the  lessening  or  annihilation  of  the 
bright  colors  of  the  males,  owing  to  their  continual  crossing  with 
the  duller  females.  It  would  be  tedious  to  follow  out  all  the 
other  possible  results;  but  I  may  remind  the  reader  that  if  sex- 
ually-limited variations  in  brightness  occurred  in  the  females, 
even  if  they  were  not  in  the  least  injurious  to  them  and  conse- 
quently were  not  eliminated,  yet  they  would  not  be  favored  or 
selected,  for  the  male  usually  accepts  any  female,  and  does  not 
select  the  more  a,ttractive  individuals;  consequently  these  varia- 
tions would  be  liable  to  be  lost,  and  would  have  little  influence 
on  the  character  of  the  race;  and  this  will  aid  in  accounting  for 
the  females  being  commonly  duller-colored  than  the  males. 

In  the  eighth  chapter  instances  were  given,  to  which  many 
might  here  be  added,  of  variations  occurring  at  various  ages,  and 
inherited  at  the  corresponding  age.  It  was  also  shown  that  va- 
riations which  occur  late  in  life  are  commonly  transmitted  to  the 
same  sex  in  which  they  first  appear;  whilst  variations  occurring 
early  in  life  are  apt  to  be  transmitted  to  both  sexes;    not  that  all 


BIRDS— DEVELOPMENT  OF  SPURS.  443 

the  cases  of  sexually-limited  transmission  can  thus  be  accounted 
for.  It  was  further  shown  that  if  a  male  bird  varied  by  becoming 
brighter  whilst  young,  such  variations  would  be  of  no  service 
until  the  age  for  reproduction  had  arrived,  and  there  was  compe- 
tition between  rival  males.  But  in  the  case  of  birds  living  on  the 
ground  and  commonly  in  need  of  the  protection  of  dull  colors, 
bright  tints  would  be  far  more  dangerous  to  the  young  and  in- 
experienced, than  to  the  adult  males.  Consequently  the  males 
which  varied  in  brightness  whilst  young  would  suffer  much  de- 
struction and  be  eliminated  through  natural  selection;  on  the 
other  hand,  the  males  which  varied  in  this  manner  when  nearly 
mature,  notwithstanding  that  they  were  exposed  to  some  addi- 
tional danger,  might  survive,  and  from  being  favored  through 
sexual  selection,  would  procreate  their  kind.  As  a  relation  often 
exists  between  the  period  of  variation  and  the  form  of  transmis- 
sion, if  the  bright-colored  young  males  were  destroyed  and  the 
mature  ones  were  successful  in  their  courtship,  the  males  alone 
would  acquire  brilliant  colors  and  would  transmit  them  exclusive- 
ly to  their  male  offspring.  But  I  by  no  means  wish  to  maintain 
that  the  influence  of  age  on  the  form  of  transmission,  is  the  sole 
cause  of  the  great  difference  in  brilliancy  between  the  sexes  of 
many  birds. 

When  the  sexes  of  birds  differ  in  color,  it  is  interesting  to 
determine  whether  the  males  alone  have  been  modified  by  sexual 
selection,  the  females  having  been  left  unchanged,  or  only  par- 
tially and  indirectly  thus  changed;  or  whether  the  females  have 
been  specially  modified  through  natural  selection  for  the  sake  of 
protection.  I  will  therefore  discuss  this  question  at  some  length, 
even  more  fully  than  its  intrinsic  importance  deserves;  for  va- 
rious curious  collateral  points  may  thus  be  conveniently  con- 
sidered. 

Before  we  enter  on  the  subject  of  color,  more  especially  in 
reference  to  Mr.  Wallace's  conclusions,  it  may  be  useful  to  dis- 
cuss some  other  sexual  differences  under  a  similar  point  of 
view.  A  breed  of  fowls  formerly  existed  in  Germany''  in  which 
the  hens  were  furnished  with  spurs;  they  were  good  layers,  but 
they  so  greatly  disturbed  their  nests  with  their  spurs  that  they 
could  not  be  allowed  to  sit  on  their  own  eggs.  Hence  at  one  time 
it  appeared  to  me  prohable  that  with  the  females  of  the  wild 
Gallinacese  the  development  of  spurs  had  been  checked  through 
natural  selection,  from  the  injury  thus  caused  to  their  nests. 
This  seemed  all  the  more  probable,  as  wing-spurs,  which  would 
not  be  injurious  during  incubation,  are  often  as  well-developed 
in  the  female  as  in  the  male;    though  in  not  a  few  cases  they  are 


•  Bechstein,  'Naturgesch.  Deutschlands,'  1793,  B.  iii.  s.  339, 


444  THE  DESCENT  OF  MAN. 

rather  larger  in  tlie  male.  When  the  male  is  furnished  with 
leg-spurs  the  female  almost  always  exhibits  rudiments  of  them, 
— the  rudiment  sometimes  consisting  of  a  mere  scale,  as  in  Gallus. 
Hence  it  might  be  argued  that  the  females  had  aboriginally  been 
furnished  with  well-developed  spurs,  but  that  these  had  subse- 
quently been  lost  through  disuse  or  natural  selection.  But  if  this 
view  be  admitted,  it  would  have  to  be  extended  to  innumerable 
other  cases;  and  it  implies  that  the  female  progenitors  of  the  ex- 
isting spur-bearing  species  were  once  encumbered  with  an  in- 
jurious appendage. 

In  some  few  genera  and  species,  as  in  Galloperdix,  Acomus,  and 
the  Javan  peacock  (Pavo  muticus),  the  females,  as  well  as  the 
males,  possess  well-developed  leg-spurs.  Are  we  to  infer  from 
this  fact,  that  they  construct  a  different  sort  of  nest  from  that 
made  by  their  nearest  allies,  and  not  liable  to  be  injured  by  their 
spurs;  so  that  the  spurs  have  not  been  removed.  Or  are  we  to 
suppose  that  the  females  of  these  several  species  especially  re- 
quire spurs  for  their  defense?  It  is  a  more  probable  conclusion 
that  both  the  presence  and  absence  of  spurs  in  the  females  result 
from  different  laws  of  inheritance  having  prevailed,  independent- 
ly of  natural  selection.  With  the  many  females  in  which  spurs 
appear  as  rudiments,  we  may  conclude  that  some  few  of  the 
successive  variations,  through  which  they  were  developed  in  the 
males,  occurred  very  early  in  life,  and  were  consequently  trans- 
ferred to  the  females.  In  the  other  and  much  rarer  cases,  in 
which  the  females  possess  fully  developed  spurs,  we  may  conclude 
that  all  the  successive  variations  were  transferred  to  them;  and 
that  they  gradually  acquired  and  inherited  the  habit  of  not  dis- 
turbing their  nests. 

The  vocal  organs  and  the  feathers  variously-modified  for  pro- 
ducing sound,  as  well  as  the  proper  instincts  for  using  them, 
often  differ  in  the  two  sexes,  but  are  sometimes  the  same  in  both. 
Can  such  differences  be  accounted  for  by  the  males  having  ac- 
quired these  organs  and  instincts,  whilst  the  females  have  been 
saved  from  inheriting  them,  on  account  of  the  danger  to  which 
they  would  have  been  exposed  by  attracting  the  attention  of  birds 
or  beasts  of  prey?  This  does  not  seem  to  me  probable,  when  we 
think  of  the  multitude  of  birds  which  with  impunity  gladden  the 
country  with  their  voices  during  the  spring.^  It  is  a  safer  con- 
clusion that,  as  vocal  and  instrumental  organs  are  of  special  ser- 
vice only  to  the  males  during  their  courtship,  these  organs  were 

'  Daines  Barrington,  however,  thought  it  probable  ('Phil.  Transact.' 
1773,  p.  164)  that  few  female  birds  sing,  because  the  talent  would  have 
been  dangerous  to  them  during  incubation.  He  adds,  that  a  similar 
view  may  possibly  account  for  the  inferiority  of  the  female  to  the  male 
in  plumage. 


BIRDS— LENGTH   OF   FEMALE'S   TAIL.  445 

developed  through  sexual  selection  and  their  constant  use  in 
that  sex  alone — the  successive  variations  and  the  effects  of  use 
having  been  from  the  first  more  or  less  limited  in  transmission 
to  the  male  offspring. 

Many  analogous  cases  could  be  adduced;  those  for  instance  of 
the  plumes  on  the  head  being  generally  longer  in  the  male  than 
in  the  female,  sometimes  of  equal  length  in  both  sexes,  and  oc- 
casionally absent  in  the  female, — these  several  cases  occurring  in 
the  same  group  of  birds.  It  would  be  diflBcult  to  account  for  such 
a  difference  between  the  sexes  by  the  female  having  been  bene- 
fited by  possessing  a  slightly  shorter  crest  than  the  male,  and 
its  consequent  diminution  or  complete  suppression  through  natu- 
ral selection.  But  I  will  take  a  more  favorable  case,  namely  the 
length  of  the  tail.  The  long  train  of  the  peacock  would  have 
been  not  only  inconvenient  but  dangerous  to  the  peahen  during 
the  period  of  incubation  and  whilst  accompanying  her  young. 
Hence  there  is  not  the  least  a  priori  improbability  in  the  de- 
velopment of  her  tail  having  been  checked  through  natural  selec- 
tion. But  the  females  of  various  pheasants,  which  apparently 
are  exposed  on  their  open  nests  to  as  much  danger  as  the  peahen, 
have  tails  of  considerable  length.  The  females  as  well  as  the 
males  of  the  Menura  superba  have  long  lails,  and  they  build  a 
domed  nest,  which  is  a  great  anomaly  in  so  large  a  bird.  Natu- 
ralists have  v/ondered  how  the  female  Menura  could  manage  her 
tail  during  incubation;  but  it  is  now  known^  that  she  "enters  the 
"nest  head  first,  and  then  turns  round  with  her  tail  sometimes  over 
"her  back,  but  more  often  bent  round  by  her  side.  Thus  in  time 
"the  tail  becomes  quite  askew,  and  is  a  tolerable  guide  to  the 
"length  of  time  the  bird  has  been  sitting."  Both  sexes  of  an 
Australian  kingfisher  (Tanysiptera  sylvia)  have  the  middle  tail- 
feathers  greatly  lengthened,  and  the  female  makes  her  nest  in  a 
hole;  and  as  I  am  informed  by  Mr.  R.  B.  Sharpe  these  feathers 
become  much  crumpled  during  incubation. 

In  these  two  latter  cases  the  great  length  of  the  tail-feathers 
must  be  in  some  degree  inconvenient  to  the  female;  and  as  in 
both  species  the  tail-feathers  of  the  female  are  somewhat  shorter 
than  those  of  the  male,  it  might  be  argued  that  their  full  develop- 
ment had  been  prevented  through  natural  selection.  But  if  the 
development  of  the  tail  of  the  peahen  had  been  checked  only 
when  it  became  inconveniently  or  dangerously  great,  she  would 
have  retained  a  much  longer  tail  than  she  actually  possesses; 
for  her  tail  is  not  nearly  so  long,  relatively  to  the  size  of  her 
body,  as  that  of  many  female  pheasants,  nor  longer  than  that 
of  the  female  turkey.  It  must  also  be  borne  in  mind,  that  in 
accordance  with  this  view  as  soon  as  the  tail  of  the  peahen  he- 

^  Mr.  Ramsay,  in  'Proc.  Zoolog-.  Soc'  1868,  p.  50. 


44«  THE  DESCENT  OF  MAN. 

came  dangerously  long,  and  its  development  was  consequently 
checked,  she  would  have  continually  reacted  on  her  male  progeny, 
and  thus  have  prevented  the  peacock  from  acquiring  his  present 
magnificent  train.  We  may  therefore  infer  that  the  length  of 
the  tail  in  the  peacock  and  its  shortness  in  the  peahen  are  the 
result  of  the  requisite  variations  in  the  male  having  been  from 
the  first  transmitted  to  the  male  offspring  alone. 

We  are  led  to  a  nearly  similar  conclusion  with  respect  to  the 
length  of  the  tail  in  the  various  species  of  pheasants.  In  the 
Eared  pheasant  (Crossoptilon  auritum)  the  tail  is  of  equal  length 
in  both  sexes,  namely,  sixteen  or  seventeen  inches;  in  the  com- 
mon pheasant  it  is  about  twenty  inches  long  in  the  male  and 
twelve  in  the  female;  in  Scemmerring's  pheasant,  thirty-seven 
Inches  in  the  male  and  only  eight  in  the  female;  and  lastly  in 
Reeve's  pheasant  it  is  sometimes  actually  seventy-two  inches 
long  in  the  male  and  sixteen  in  the  female.  Thus  in  the  several 
species,  the  tail  of  the  female  differs  much  in  length,  irrespective- 
ly of  that  of  the  male;  and  this  can  be  accounted  for,  as  it  seems 
to  me,  with  much  more  probability,  by  the  laws  of  inheritance, — 
that  is  by  the  successive  variations  having  been  from  the  first 
more  or  less  closely  limited  in  their  transmission  to  the  male  sex, 
than  by  the  agency  of  natural  selection,  resulting  from  the  length 
of  tail  being  more  or  less  injurious  to  the  females  of  these  several 
allied  species. 

We  may  now  consider  Mr.  Wallace's  arguments  in  regard  to 
the  sexual  coloration  of  birds.  He  believes  that  the  bright  tints 
originally  acquired  through  sexual  selection  by  the  males,  would 
in  all,  or  almost  all  cases,  have  been  transmitted  to  the  females, 
unless  the  transference  had  been  checked  through  natural  selec- 
tion. I  may  here  remind  the  reader  that  various  facts  opposed 
to  this  view  have  already  been  given  under  reptiles,  amphibians, 
fishes,  and  lepidoptera.  Mr.  Wallace  rests  his  belief  chiefly,  but 
not  exclusively,  as  we  shall  see  in  the  next  chapter,  on  the  fol- 
lowing statement,^  that  when  both  sexes  are  colored  in  a  very 
conspicuous  manner,  the  nest  is  of  such  a  nature  as  to  conceal 
the  sitting  bird;  but  when  there  is  a  marked  contrast  of  color 
between  the  sexes,  the  male  being  gay  and  the  female  dull-col- 
ored, the  nest  is  open  and  exposes  the  sitting  bird  to  view.  This 
coincidence,  as  far  as  it  goes,  certainly  seems  to  favor  the  belief 
that  the  females  which  sit  on  open  nests  have  been  specially  modi- 
fied for  the  sake  of  protection;  but  we  shall  presently  see  that 
there  is  another  and  more  probable  explanation,  namely,  that 
conspicuous  females  have  acquired  the  instinct  of  building  domed 
nests  oftener  than  dull-colored  birds.    Mr.  Wallace  admits  that 

•  'Journal  of  Travel/  edited  by  A.  Murray,  vol.  i.  1868,  p.  78. 


BIRDS— COLOR   AND    NIDIFICATION.  447 

there  are,  as  might  have  been  expected,  some  exceptions  to  his 
two  rules,  but  it  is  a  question  whether  the  exceptions  are  not  so 
numerous  as  seriously  to  invalidate  them. 

There  is  in  the  first  place  much  truth  in  the  Duke  of  Argyll's 
remark^°  that  a  large  domed  nest  is  more  conspicuous  to  an  enemy, 
especially  to  all  tree-haunting  carnivorous  animals,  than  a  smaller 
open  nest.  Nor  must  we  forget  that  with  many  birds  which  build 
open  nests,  the  male  sits  on  the  eggs  and  aids  the  female  in 
feeding  the  young:  this  is  the  case,  for  instance,  with  Pyranga 
asstiva,^^  one  of  the  most  splendid  birds  in  the  United  States,  the 
male  being  vermilion,  and  the  female  light  brownish-green.  Now 
if  brilliant  colors  had  been  extremely  dangerous  to  birds  whilst 
sitting  on  their  open  nests,  the  males  in  these  cases  would  have 
suffered  greatly.  It  might,  however,  be  of  such  paramount  im- 
portance to  the  male  to  be  brilliantly  colored,  in  order  to  beat 
his  rivals,  that  this  may  have  more  than  compensated  some  addi- 
tional danger. 

Mr.  Wallace  admits  that  with  the  King-crows  (Dicrurus),  Ori- 
oles, and  Pittidse,  the  females  are  conspicuously  colored,  yet 
build  open  nests;  but  he  urges  that  the  birds  of  the  first  group 
are  highly  pugnacious  and  could  defend  themselves;  that  those 
of  the  second  group  take  extreme  care  in  concealing  their  open 
nests,  but  this  does  not  invariably  hold  good;^-  and  that  with  the 
birds  of  the  third  group  the  females  are  brightly  colored  chiefly 
on  the  under  surface.  Besides  these  cases,  pigeons  which  are 
sometimes  brightly,  and  almost  always  conspicuously  colored, 
and  which  are  notoriously  liable  to  the  attacks  of  birds  of  prey, 
offer  a  serious  exception  to  the  rule,  for  they  almost  always  build 
open  and  exposed  nests.  In  another  large  family,  that  of  the 
humming-birds,  all  the  species  build  open  nests,  yet  with  some  of 
the  most  gorgeous  species  the  sexes  are  alike;  and  in  the  major- 
ity, the  females,  though  less  brilliant  than  the  males,  are  brightly 
colored.  Nor  can  it  be  maintained  that  all  female  humming- 
birds, which  are  brightly  colored,  escape  detection  by  their  tints 
being  green,  for  some  display  on  their  upper  surfaces  red,  blue, 
and  other  colors.^" 


10  'Journal  of  Travel,'  edited  by  A.  Murray,  vol.  i.  1S68,  p.  281. 

iiAudubon,  'Ornithological  Biography,'  vol.  i.  p.  233. 

isjerdon,  'Birds  of  India,'  vol.  ii.  p.  108.  Gould's  'Handbook  of  the 
Birds  of  Australia,'  vol.  i.  p.  463. 

^  For  instance,  the  female  Eupetomena  macroura  has  the  head  and 
tail  dark  blue  with  reddish  loins;  the  female  Lampornis  porphyrurus 
is  blackish-green  on  the  upper  surface,  with  the  lores  and  sides  of  the 
throat  crimson;  the  female  Eulampis  jugularis  has  the  top  of  the 
head  and  back  green,  but  the  loins  and  the  tail  are  crimson.  Many 
other  instances  of  highly  conspicuous  females  could  be  given.  See 
Mr.  Gould's  magnificent  work  on  this  family. 


448  THE  DESCENT  OF  MAN. 

In  regard  to  birds  whicli  build  in  holes  or  construct  domed 
nests,  other  advantages,  as  Mr.  Wallace  remarks,  besides  con- 
cealment are  gained,  such  as  shelter  from  the  rain,  greater 
warmth,  and  in  hot  countries  protection  from  the  sun;^*  so  that 
it  is  no  valid  objection  to  his  view  that  many  birds  having  both 
sexes  obscurely  colored  build  concealed  nests.'^  The  female  Horn- 
bill  (Buceros),  for  instance,  of  India  and  Africa  is  protected  dur- 
ing incubation  with  extraordinary  care,  for  she  plasters  up  with 
her  own  excrement  the  orifice  of  the  hole  in  which  she  sits  on  her 
eggs,  leaving  only  a  small  orifice  through  which  the  male  feeds 
her;  she  is  thus  kept  a  close  prisoner  during  the  whole  period 
of  incubation  ;^*'  yet  female  horn-bills  are  not  more  conspicuously 
colored  than  many  other  birds  of  equal  size  which  build  open 
nests.  It  is  a  more  serious  objection  to  Mr.  Wallace's  view,  as  is 
admitted  by  him,  that  in  some  few  groups  the  males  are  bril- 
liantly colored  and  the  females  obscure,  and  yet  the  latter  hatch 
their  eggs  in  domed  nests.  This  is  the  case  with  the  GrallinaB 
of  Australia,  the  Superb  Warblers  (Maluridse)  of  the  same  coun- 
try, the  Sun-birds  (Nectariniae),  and  with  several  of  the  Aus- 
tralian Honey-suckers  or  Meliphagidae." 

If  we  look  to  the  birds  of  England  we  shall  see  that  there  is  no 
close  and  general  relation  between  the  colors  of  the  female  and 
the  nature  of  the  nest  which  is  constructed.  About  forty  of  our 
British  birds  (excluding  those  of  large  size  which  could  defend 
themselves)  build  in  holes  in  banks,  rocks,  or  trees,  or  construct 
domed  nests.  If  we  take  the  colors  of  the  female  goldfinch,  bull- 
finch, or  blackbird,  as  a  standard  of  the  degree  of  conspicuousness, 
which  is  not  highly  dangerous  to  the  sitting  female,  then  out 
of  the  above  forty  birds,  the  females  of  only  twelve  can  be  con- 
sidered as  conspicuous  to  a  dangerous  degree,  the  remaining 
twenty-eight  being  inconspicuous.^®    Nor  is  there  any  close  rela- 

1*  Mr.  SaJvin  noticed  in  Guatemala  ('Ibis,'  1864,  p.  375)  that  humming- 
birds were  much  more  unwilling  to  leave  their  nests  during  very  hot 
weather,  when  the  sun  was  shining  brightly,  as  if  their  eggs  would 
be  thus  injured,  than  during  cool,  cloudy,  or  rainy  weather. 

13 1  may  specify,  as  instances  of  dull-colored  birds  building  concealed 
nests,  the  species  belonging  to  eight  Australian  g'enera,  described  in 
Gould's  'Handbook  of  the  Birds  of  Australia,'  vol.  i.  pp.  340,  362,  365, 
383,  387,  389,  391,  414. 

18  Mr.  C.  Home,  'Proc.  Zoolog.  Soc'  1869,  p.  243. 

"  On  the  nidification  and  colors  of  these  latter  species,  see  Gould's 
'Handbook,'  &c.,  vol.  i.  pp.  504,  527. 

18  I  have  consulted  on  this  subject,  Macgillivray's  'British  Birds,' 
and  though  doubts  may  be  entertained  in  some  cases  in  regard  to 
the  degree  of  concealment  of  the  nest,  and  to  the  degree  of  con- 
spicuousness of  the  female,  yet  the  following  birds,  which  all  lay 
their  eggs  in  holes  or  in  domed  nests,  can  hardly  be  considered,  by 


BIRDS— COLOR   AND    NIDIFICATION.  449 

tion  within  the  same  genus  between  a  well-pronounced  difference 
in  color  between  the  sexes,  and  the  nature  of  the  nest  constructed. 
Thus  the  male  house  sparrow  (Passer  domesticus)  differs  much 
from  the  female,  the  male  tree-sparrow  (P.  montanus)  hardly  at 
all,  and  yet  both  build  well-concealed  nests.  The  two  sexes  of  the 
common  fly-catcher  (Musicapse  grisola)  can  hardly  be  distin- 
guished, whilst  the  sexes  of  the  pied  fly-catcher  (M.  luctuosa)  dif- 
fer considerably,  and  both  species  build  in  holes  or  conceal  their 
nests.  The  female  blackbird  (Turdus  merula)  differs  much,  the 
female  ring-ouzel  (T.  torquatus)  differs  less,  and  the  female  com- 
mon thrush  (T.  musicus)  hardly  at  all  from  their  respective  males; 
yet  all  build  open  nests.  On  the  other  hand,  the  not  very  dis- 
tantly-allied water-ouzel  (Cinclus  aquaticus)  builds  a  domed  nest, 
and  the  sexes  differ  about  as  much  as  in  the  ring-ouzel.  The. 
black  and  red  grouse  (Tetrao  tetrix  and  T.  scoticus)  build  open 
nests  in  equally  well-concealed  spots,  but  in  the  one  species  the 
sexes  differ  greatly,  and  in  the  other  very  little. 

Notwithstanding  the  foregoing  objections,  I  cannot  doubt,  after 
reading  Mr.  Wallace's  excellent  essay,  that  looking  to  the  birds 
of  the  world,  a  large  majority  of  the  species  in  which  the  females 
are  conspicuously  colored  (and  in  this  case  the  males  with  rare 
exceptions  are  equally  conspicuous),  build  concealed  nests  for 
the  sake  of  protection.  Mr.  Wallace  enumerates^''  a  long  series 
of  groups  in  which  this  rule  holds  good;  but  it  will  suffice  here 
to  give,  for  instance,  the  more  familiar  groups  of  kingfishers, 
toucans,  trogons,  puff-birds  (Capitonid^),  plantain-eaters  (Mu- 
sophagse),  woodpeckers,  and  parrots.  Mr.  Wallace  believes  that 
in  these  groups,  as  the  males  gradually  acquired  through  sexual 
selection  their  brilliant  colors,  these  were  transferred  to  the  fe- 
males and  were  not  eliminated  by  natural  selection,  owing  to 
the  protection  which  they  already  enjoyed  from  their  manner  of 
nidification.  According  to  this  view,  their  present  manner  of 
nesting  was  acquired  before  their  present  colors.  But  it  seems  to 
me  much  more  probable  that  in  most  cases,  as  the  females  were 
gradually  rendered  more  and  more  brilliant  from  partaking  of 
the  colors  of  the  male,  they  were  gradually  led  to  change  their  in- 
stincts (supposing  that  they  originally  built  open  nests),  and  to 


the  above  standard,  as  conspicuous:  Passer,  2  species;  Sturnus,  of 
which  the  female  is  considerably  less  briUiant  than  the  male;  Cinclus; 
Motacilla  boarula  (?);  Erithacus  (?);  Fruticola,  2  sp.;  Saxicola;  Ruti- 
cilla,  2  sp.;  Sylvia,  3  sp.;  Parus,  3  sp.;  Mecistura;  Anorthura;  Cer- 
thia;  Sitta;  Yunx;  Muscieapa,  2  sp. ;  Hirundo,  3  sp.;  and  Cypselus. 
The  females  of  the  following  12  birds  may  be  considered  as  conspicu- 
ous, according  to  the  same  standard,  viz.,  Pastor,  Motacilla  alba, 
Parus  major  and  P.  caeruleus,  Upupa,  Picus,  4  sp.,  Coracias,  Alcedo, 
and  Merops. 
18  'Journal  of  Travel,'  edited  by  A.  Murray,  vol.  i.  p.  78. 
30 


450  THE  DESCENT  OF  MAN. 

seek  protection  by  building  domed  or  concealed  nests.  No  one 
who  studies,  for  instance,  Audubon's  account  of  the  differences  in 
the  nests  of  the  same  species  in  the  Northern  and  Southern  United 
States,-"  will  feel  any  great  difficulty  in  admitting  that  birds, 
either  by  a  change  (in  the  strict  sense  of  the  word)  of  their  habits, 
or  through  the  natural  selection  of  so-called  spontaneous  varia- 
tions of  instinct,  might  readily  be  led  to  modify  their  manner  of 
nesting. 

This  way  of  viewing  the  relation,  as  far  as  it  holds  good,  be- 
tween the  bright  colors  of  female  birds  and  their  manner  of 
nesting,  receives  some  support  from  certain  cases  occurring  in 
the  Sahara  Desert.  Here,  as  in  most  other  deserts,  various  birds, 
and  many  other  animals,  have  had  their  colors  adapted  in  a 
wonderful  manner  to  the  tints  of  the  surrounding  surface.  Never- 
theless there  are,  as  I  am  informed  by  the  Rev.  Mr.  Tristram,  some 
curious  exceptions  to  the  rule;  thus  the  male  of  the  Monticola 
cyanea  is  conspicuous  from  his  bright  blue  color,  and  the  female 
almost  equally  conspicuous  from  her  mottled  brown  and  white 
plumage;  both  sexes  of  two  species  of  Dromolsea  are  of  a  lustrous 
black;  so  that  these  three  species  are  far  from  receiving  protec- 
tion from  their  colors,  yet  they  are  able  to  survive,  for  they  have 
acquired  the  habit  of  taking  refuge  from  danger  in  holes  or 
crevices  in  the  rocks. 

With  respect  to  the  above  groups  in  which  the  females  are 
conspicuously  colored  and  build  concealed  nests,  it  is  not  neces- 
sary to  suppose  that  each  separate  species  had  its  nidifying  in- 
stinct specially  modified;  but  only  that  the  early  progenitors  of 
each  group  were  gradually  led  to  build  domed  or  concealed  nests, 
and  afterwards  transmitted  this  instinct,  together  with  their 
bright  colors,  to  their  modified  descendants.  As  far  as  it  can  be 
trusted,  the  conclusion  is  interesting,  that  sexual  selection,  to- 
gether with  equal  or  nearly  equal  inheritance  by  both  sexes,  have 
indirectly  determined  the  manner  of  nidification  of  whole  groups 
of  birds. 

According  to  Mr.  Wallace,  even  in  the  groups  in  which  the 
females,  from  being  protected  in  domed  nests  during  incubation, 
have  not  had  their  bright  colors  eliminated  through  natural 
selection,  the  males  often  differ  in  a  slight,  and  occasionally  in  a 
considerable  degree,  from  the  females.  This  is  a  significant  fact, 
for  such  differences  in  color  must  be  accounted  for  by  some  of 
the  variations  in  the  males  having  been  from  the  first  limited  in 
transmission  to  the  same  sex;  as  it  can  hardly  be  maintained  that 
these  differences,  especially  when  very  slight,  serve  as  a  protec- 

20  See  many  statements  in  the  'Ornithological  Biography.'  See,  also, 
some  curious  observations  on  the  nests  of  Italian  birds  by  Eugenio  Bet- 
toni,  in  the  'Atti  della  Societa  Italiana,'  vol.  xi.  1869,  p.  487. 


BIRDS— COLOR   AND    NIDIFICATION.  451 

tion  to  the  female.  Thus  all  the  species  in  the  splendid  group 
of  the  Trogons  build  in  holes;  and  Mr.  Gould  gives  figures-^  of 
both  sexes  of  twenty-five  species,  in  all  of  which,  with  one  partial 
exception,  the  sexes  differ  sometimes  slightly,  sometimes  con- 
spicuously, in  color,— the  males  being  always  finer  than  the  fe- 
males, though  the  latter  are  likewise  beautiful.  All  the  species 
of  kingfishers  build  in  holes,  and  with  most  of  the  species  the 
sexes  are  equally  brilliant,  and  thus  far  Mr.  Wallace's  rule  holds 
good;  but  in  some  of  the  Australian  species  the  colors  of  the 
females  are  rather  less  vivid  than  those  of  the  male;  and  in  one 
splendidly-colored  species,  the  sexes  differ  so  much  that  they 
were  at  first  thought  to  be  specifically  distinct.-"  Mr.  R.  B.  Sharpe, 
who  has  especially  studied  this  group,  has  shown  me  some  Amer- 
ican species  (Ceryle)  in  which  the  breast  of  the  male  is  belted 
with  black.  Again,  in  Carcineutes,  the  difference  between  the 
sexes  is  conspicuous:  in  the  male  the  upper  surface  is  dull-blue 
banded  with  black,  the  lower  surface  being  partly  fawn-colored, 
and  there  is  much  red  about  the  head;  in  the  female  the  upper 
surface  is  reddish-brown  banded  with  black,  and  the  lower  sur- 
face white  with  black  markings.  It  is  an  interesting  fact,  as 
showing  how  the  same  peculiar  style  of  sexual  coloring  often 
characterizes  allied  forms,  that  in  three  species  of  Dacelo  the 
male  differs  from  the  female  only  in  the  tail  being  dull-blue  banded 
with  black,  whilst  that  of  the  female  is  brown  with  blackish 
bars;  so  that  here  the  tail  differs  in  color  in  the  two  sexes  in 
exactly  the  same  manner  as  the  whole  upper  surface  in  the  two 
sexes  of  Carcineutes. 

With  parrots,  which  likewise  build  in  holes,  we  find  analogous 
cases:  in  most  of  these  species  both  sexes  are  brilliantly  colored 
and  indistinguishable,  but  in  not  a  few  species  the  males  are 
colored  rather  more  vividly  than  the  females,  or  even  very  differ- 
ently from  them.  Thus,  besides  other  strongly-marked  differ- 
ences, the  whole  under  surface  of  the  male  King  Lory  (Apros- 
mictus  scapulatus)  is  scarlet,  whilst  the  throat  and  chest  of  the 
female  is  green  tinged  with  red:  in  the  Buphema  splendida  there 
is  a  similar  difference,  the  face  and  wing-coverts  moreover  of 
the  female  being  of  a  paler  blue  than  in  the  male."^  In  the  family 
of  the  tits  (Parinse),  which  build  concealed  nests,  the  female  of 
our  common  blue  tomtit  (Parus  cseruleus)  is  "much  less  brightly 

81  See  his  'Monograph  of  the  Trogonidae,'  first  edition. 

22  Namely  Cyanalcyon.  Gould's  'Handbook  to  the  Birds  of  Australia/ 
vol.  i.  p.  133;    see,  also,  pp.  130,  136. 

23  Every  gradation  of  difference  between  the  sexes  may  be  followed 
in  the  parrots  of  Australia.  See  Gould's  'Handbook'  &c.,  vol.  ii. 
pp.  14-102. 


452  THE  DESCENT  OF  MAN. 

"colored"  than  the  male;    and  in  the  magnificent  Sultan  yellow  tit 
of  India  the  difference  is  greater.-* 

Again  in  the  great  army  of  the  woodpeckers,-^  the  sexes  are 
generally  nearly  alike,  but  in  the  Megapicus  validus  all  those 
parts  of  the  head,  neck,  and  breast,  which  are  crimson  in  the 
male  are  pale  brown  in  the  female.  As  in  several  woodpeckers 
the  head  of  the  male  is  bright  crimson,  whilst  that  of  the  female 
is  plain,  it  occurred  to  me  that  this  color  might  possibly  make 
the  female  dangerously  conspicuous,  whenever  she  put  her  head 
out  of  the  hole  containing  her  nest,  and  consequently  that  this 
color,  in  accordance  with  Mr.  Wallace's  belief,  had  been  elim- 
inated. This  view  is  strengthened  by  what  Malherbe  states  with 
respect  to  Indopicus  carlotta;  namely,  that  the  young  females, 
like  the  young  males,  have  some  crimson  about  their  heads,  but 
that  this  color  disappears  in  the  adult  female,  whilst  it  is  intent 
sified  in  the  adult  male.  Nevertheless  the  following  considera- 
tions render  this  view  extremely  doubtful:  the  male  takes  a 
fair  share  in  incubation,^"  and  would  be  thus  almost  equally  ex- 
posed to  danger;  both  sexes  of  many  species  have  their  heads 
of  an  equally  bright  crimson;  in  other  species  the  difference  be- 
tween the  sexes  in  the  amount  of  scarlet  is  so  slight  that  it  can 
hardly  make  any  appreciable  difference  in  the  danger  incurred; 
and  lastly,  the  coloring  of  the  head  in  the  two  sexes  often  differs 
slightly  in  other  ways. 

The  cases,  as  yet  given,  of  slight  and  graduated  differences  in 
color  between  the  males  and  females  in  the  groups,  in  which  as 
a  general  rule  the  sexes  resemble  each  other,  all  relate  to  species 
which  "build  domed  or  concealed  nests.  But  similar  gradations 
may  likewise  be  observed  in  groups  in  which  the  sexes  as  a 
general  rule  resemble  each  other,  but  which  build  open  nests. 
As  I  have  before  instanced  the  Australian  parrots,  so  I  may  here 
instance,  without  giving  any  details,  the  Australian  pigeons." 
It  deserves  especial  notice  that  in  all  these  cases  the  slight  dif- 
ferences in  plumage  between  the  sexes  are  of  the  same  general 
nature  as  the  occasionally  greater  differences.  A  good  illustra- 
tion of  this  fact  has  already  been  afforded  by  those  kingfishers  in 
which  either  the  tail  alone  or  the  whole  upper  surface  of  the 
plumage  differs  in  the  same  manner  in  the  two  sexes.  Similar 
cases  may  be  observed  with  parrots  and  pigeons.    The  differences 


2*  Macgillivray's  'British  Birds,'  vol.  ii.  p.  433.  Jerdon,  'Birds  of  In- 
dia,' vol.  ii.  p.  282. 

-5  All  the  following  facts  are  taken  from  M.  Malherbe' s  magnificent 
'Monographic  des  Picidees,'  1861. 

28  Audubon's  'Ornithological  Biography,'  vol.  ii.  p.  75;  see,  also,  the 
'Ibis,'  vol.  1.  p.  268. 

27  Gould's  'Handbook  to  the  Birds  of  Australia,'  vol.  ii.  pp.  109-149. 


BIRDS— COLOR   AND    NIDIFICATION.  453 

in  color  between  the  sexes  of  the  same  species  are,  also,  of  the 
same  general  nature  as  the  differences  in  color  between  the  dis- 
tinct species  of  the  same  group.  For  when  in  a  group  in  which 
the  sexes  are  usually  alike,  the  male  differs  considerably  from 
the  female,  he  is  not  colored  in  a  quite  new  style.  Hence  we 
may  infer  that  within  the  same  group  the  special  colors  of  both 
sexes  when  they  are  alike,  and  the  colors  of  the  male,  when  he 
differs  slightly  or  even  considerably  from  the  female,  have  been  in 
most  cases  determined  by  the  same  general  cause;  this  being 
sexual  selection. 

It  is  not  probable,  as  has  already  been  remarked,  that  differ- 
ences in  color  between  the  sexes,  when  very  slight,  can  be  of 
service  to  the  female  as  a  protection.  Assuming,  however,  that 
they  are  of  service,  they  might  be  thought  to  be  cases  of  transi- 
tion; but  we  have  no  reason  to  believe  that  many  species  at 
any  one  time  are  undergoing  change.  Therefore  we  can  hardly 
admit  that  the  numerous  females  which  differ  very  slightly  in 
color  from  the'.r  males  are  now  all  commencing  to  become  ob- 
scure for  the  sake  of  protection.  Even  if  we  consider  somewhat 
more  marked  sexual  differences,  is  it  probable,  for  instance,  that 
the  head  of  the  female  chaffinch, — the  crimson  on  the  breast  of 
the  female  bullfinch, — the  green  of  the  female  greenfinch, — the 
crest  of  the  female  golden-crested  wren,  hctve  all  been  rendered 
less  bright  by  the  slow  process  of  selection  for  the  sake  of  protec- 
tion? I  cannot  think  so;  and  still  less  with  the  slight  differences 
between  the  sexes  of  these  birds  which  build  concealed  nests.  On 
the  other  hand,  the  differences  in  color  between  the  sexes,  whether 
great  or  small,  may  to  a  large  extent  be  explained  on  the  principle 
of  the  successive  variations,  acquired  by  the  males  through  sex- 
ual selection,  having  been  from  the  first  more  or  less  limited  in 
their  transmission  to  the  females.  That  the  degree  of  limitation 
should  differ  in  different  species  of  the  same  group  will  not  sur- 
prise any  one  who  has  studied  the  laws  of  inheritance,  for  they 
are  so  complex  that  they  appear  to  us  in  our  ignorance  to  be 
capricious  in  their  action.-^ 

As  far  as  I  can  discover  there  are  few  large  groups  of  birds  in 
which  all  the  species  have  both  sexes  alike  and  brilliantly  col- 
ored, but  I  hear  from  Mr.  Sclater,  that  this  appears  to  be  the  case 
with  the  Musophagse  or  plantain-eaters.  Nor  do  I  believe  that 
any  large  groups  exist  in  which  the  sexes  of  all  the  species  are 
widely  dissimilar  in  color:  Mr.  Wallace  informs  me  that  the 
chatterers  of  S.  America  (Cotingidae)  offer  one  of  the  best  in- 
stances; but  with  some  of  the  species,  in  which  the  male  has  a 
splendid  red  breast,  the  female  exhibits  some  red  on  her  breast; 

28  See  remarks  to  this  effect  in  my  work  on  'Variation  ujider  Domesti- 
cation,' vol.  ii.  chap.  xii. 
30 


454  THE  DESCENT  OF  MAN. 

and  the  females  of  other  species  show  traces  of  the  green  and 
other  colors  of  the  males.  Nevertheless  we  have  a  near  approach 
to  close  sexual  similarity  or  dissimilarity  throughout  several 
groups:  and  this,  from  what  has  just  been  said  of  the  fluctuating 
nature  of  inheritance,  is  a  somewhat  surprising  circumstance. 
But  that  the  same  laws  should  largely  prevail  with  allied  animals 
is  not  surprising.  The  domestic  fowl  has  produced  a  great  num- 
ber of  breeds  and  sub-breeds,  and  in  these  the  sexes  generally 
differ  in  plumage;  so  that  it  has  been  noticed  as  an  unusual  cir- 
cumstance when  in  certain  sub-breeds  they  resemble  each  other. 
On  the  other  hand,  the  domestic  pigeon  has  likewise  produced 
a  vast  number  of  distinct  breeds  and  sub-breeds,  and  in  these, 
with  rare  exceptions,  the  two  sexes  are  identically  alike.  There- 
fore if  other  species  of  Gallus  and  Columba  were  domesticated  and 
varied,  it  would  not  be  rash  to  predict  that  similar  rules  of  sex- 
ual similarity  and  dissimilarity,  depending  on  the  form  of  trans- 
mission, would  hold  good  in  both  cases.  In  like  manner  the  same 
form  of  transmission  has  generally  prevailed  under  nature 
throughout  the  same  groups,  although  marked  exceptions  to  this 
rule  occur.  Thus  within  the  same  family  or  even  genus,  the 
sexes  may  be  identically  alike,  or  very  different  in  color.  In- 
stances have  already  been  given  in  the  same  genus,  as  with  spar- 
rows, fly-catchers,  thrushes  and  grouse.  In  the  family  of  pheas- 
ants the  sexes  of  almost  all  the  species  are  wonderfully  dissimilar, 
but  are  quite  alike  in  the  eared  pheasant  or  Crossoptilon  auritum. 
In  two  species  of  Chloephaga,  a  genus  of  geese,  the  male  cannot 
be  distinguished  from  the  females,  except  by  size;  whilst  in  two 
others,  the  sexes  are  so  unlike  that  they  might  easily  be  mistaken 
for  distinct  species.^* 

The  laws  of  inheritance  can  alone  account  for  the  following 
cases,  in  which  the  female  acquires,  late  in  life,  certain  characters 
proper  to  the  male,  and  ultimately  comes  to  resemble  him  more 
or  less  completely.  Here  protection  can  hardly  have  come  into 
play.  Mr.  Blyth  informs  me  that  the  females  of  Oriolus  melano- 
cephalus  and  of  some  allied  species,  when  sufilciently  mature  to 
breed,  differ  considerably  in  plumage  from  the  adult  males;  but 
after  the  second  or  third  moults  they  differ  only  in  their  beaks 
having  a  slight  greenish  tinge.  In  the  dwarf  bitterns  (Ardetta), 
according  to  the  same  authority,  "the  male  acquires  his  final 
*'livery  at  the  first  moult,  the  female  not  before  the  third  or 
"fourth  moult;  in  the  meanwhile  she  presents  an  intermediate 
"garb,  which  is  ultimately  exchanged  for  the  same  livery  as  that 
"of  the  male."  So  again  the  female  Falco  peregrinus  acquires 
her  blue  plumage  more  slowly  than  the  male.  Mr.  Swinhoe  states 
that  with  one  of  the  Drongo  shrikes  (Dicrurus  macrocercus)  the 

2»  The  'Ibis,'  vol.  vi.  1864,  p.  122. 


BIRDS— COLOR   AND    NIDIFICATION.  455 

male  whilst  almost  a  nestling,  moults  his  soft  brown  plumage  and 
iDecomes  of  a  uniform  glossy  greenish-black;  but  the  female  re- 
tains for  a  long  time  the  white  striae  and  spots  on  the  axillary- 
feathers;  and  does  not  completely  assume  the  uniform  black 
color  of  the  male  for  three  years.  The  same  excellent  observer 
remarks  that  in  the  spring  of  the  second  year  the  female  spoon- 
bill (Platalea)  of  China  resembles  the  male  of  the  first  year,  and 
that  apparently  it  is  not  until  the  third  spring  that  she  acquires 
the  same  adult  plumage  as  that  possessed  by  the  male  at  a  much 
earlier  age.  The  female  Bombycilla  carolinensis  differs  very 
little  from  the  male,  but  the  appendages,  which  like  beads  of  red 
sealing-wax  ornament  the  wing-feathers,^°  are  not  developed  in 
her  so  early  in  life  as  in  the  male.  In  the  male  of  an  Indian  par- 
rakeet  (Palseornis  javanicus)  the  upper  mandible  is  coral-red  from 
his  earliest  youth,  but  in  the  female,  as  Mr.  Blyth  has  observed 
with  caged  and  wild  birds,  it  is  at  first  black  and  does  not  be- 
come red  until  the  bird  is  at  least  a  year  old,  at  which  age  the 
sexes  resemble  each  other  in  all  respects.  Both  sexes  of  the  wild 
turkey  are  ultimately  furnished  with  a  tuft  of  bristles  on  the 
breast,  but  in  two-year-old  birds  the  tuft  is  about  four  inches  long 
in  the  male  and  hardly  apparent  in  the  female;  when,  however, 
the  latter  has  reached  her  fourth  year,  it  is  from  four  to  five  inches 
in  length.^^ 

These  cases  must  not  be  confounded  with  those  where  diseased 
or  old  females  abnormally  assume  masculine  characters,  nor  with 
those  where  fertile  females,  whilst  young,  acquire  the  characters 
Gf  the  male,  through  variation  or  some  unknown  cause.^-  But  all 
these  cases  have  so  much  in  common  that  they  depend,  according 
to  the  hypothesis  of  pangenesis,  on  gemmules  derived  from  each 
part  of  the  male  being  present,  though  latent,  in  the  female; 


so  When  the  male  courts  the  female,  these  ornaments  are  vibrated, 
and  "are  shown  off  to  great  advantage,"  on  the  outstretched  wings: 
A.  Leith  Adams,   'Field  and  Forest  Rambles,'  1873,  p.  153. 

31  On  Ardetta,  Translation  of  Cuvier's  'Regne  Animal,'  by  Mr.  Blyth, 
footnote,  p.  159.  On  the  Peregrine  Falcon,  Mr.  Blyth,  in  Charlesworth's 
'Mag.  of  Nat.  Hist.'  vol.  i.  1837,  p.  304.  On  Dicrurus,  'Ibis,'  1863,  p.  44. 
On  the  Platalea,  'Ibis,'  vol.  vi.  1864,  p.  366.  On  the  Bombycilla,  Audu- 
bon's 'Ornitholog.  Biography,'  vol.  i.  p.  229.  On  the  Palaeornis,  see, 
also,  Jerdon,  'Birds  of  India,'  vol.  i.  p.  263.  On  the  wild  turkey,  Au- 
dubon, ibid.  vol.  i.  p.  15;  but  I  hear  from  Judge  Caton  that  in  Illinois 
the  female  very  rarely  acquires  a  tuft.  Analogous  cases  with  the 
females  of  Petrocossyphus  are  given  by  Mr.  R.  B.  Sharpe,  'Proc. 
Zoolog.  Soc'  1872,  p.  496. 

32  Of  these  latter  cases  Mr.  Blyth  has  recorded  (Translation  of  Cu- 
vier's 'Regne  Animal,'  p.  158)  various  instances  with  Lanius,  Ruticilla, 
Linaria,  and  Anas.  Audubon  has  also  recorded  a  similar  case  ('Ornith 
Biog.'  vol.  V.  p.  519)  with  Lyranga  aestiva. 


456  THE  DESCENT  OF  MAN. 

their  development  following  on  some  slight  change  in  the  elective 
affinities  of  her  constituent  tissues. 

A  few  words  must  be  added  on  changes  of  plumage  in  relation 
to  the  season  of  the  year.  From  reasons  formerly  assigned  there 
can  be  little  doubt  that  the  elegant  plumes,  long  pendant  feathers, 
crests,  &c.,  of  egrets,  herons,  and  many  other  birds,  which  are 
developed  and  retained  only  during  the  summer,  serve  for  orna- 
mental and  nuptial  purposes,  though  common  to  both  sexes.  The 
female  is  thus  rendered  more  conspicuous  during  the  period  of  in- 
cubation than  during  the  winter;  but  such  birds  as  herons  and 
egrets  would  be  able  to  defend  themselves.  As,  however,  plumes 
would  probably  be  inconvenient  and  certainly  of  no  use  during 
the  winter,  it  is  possible  that  the  habit  of  moulting  twice  in  the 
year  may  have  been  gradually  acquired  through  natural  selection 
for  the  sake  of  casting  off  inconvenient  ornaments  during  the  win- 
ter. But  this  view  cannot  be  extended  to  the  many  waders,  whose 
summer  and  winter  plumages  differ  very  little  in  color.  With 
defenseless  species,  in  which  both  sexes,  or  the  males  alone,  be- 
come extremely  conspicuous  during  the  breeding-season, — or  when 
the  males  acquire  at  this  season  such  long  wing  or  tail-feathers 
as  to  impede  their  flight,  as  with  Cosmetornis  and  Vidua, — it  cer- 
tainly at  first  appears  highly  probable  that  the  second  moult  has 
been  gained  for  the  special  purpose  of  throwing  off  these  orna- 
ments. We  must,  however,  remember  that  manj'-  birds,  such  as 
some  of  the  Birds  of  Paradise,  the  Argus  pheasant  and  peacock,  do 
not  cast  their  plumes  during  the  winter;  and  it  can  hardly  be 
maintained  that  the  constitution  of  these  birds,  at  least  of  the  Gal- 
linacese,  renders  a  double  moult  impossible,  for  the  ptarmigan 
moults  thrice  in  the  year.^^  Hence  it  must  be  considered  as  doubt- 
ful whether  the  many  species  which  moult  their  ornamental 
plumes  or  lose  their  bright  colors  during  the  winter,  have  acquired 
this  habit  on  account  of  the  inconvenience  or  danger  which  they 
would  otherwise  have  suffered. 

I  conclude,  therefore,  that  the  habit  of  moulting  twice  in  the 
year  was  in  most  or  all  cases  first  acquired  for  some  distinct 
purpose,  perhaps  for  gaining  a  warmer  winter  covering;  and  that 
variations  in  the  plumage  occurring  during  the  summer  were  ac- 
cumulated through  sexual  selection,  and  transmitted  to  the  off- 
spring at  the  same  season  of  the  year;  that  such  variations  were 
inherited  either  by  both  sexes  or  by  the  males  alone,  according  to 
the  form  of  inheritance  which  prevailed.  This  appears  more 
probable  than  that  the  species  in  all  cases  originally  tended  to  re- 
tain their  ornamental  plumage  during  the  winter,  but  were  saved 


23  See  Gould's  'Birds  of  Great  Britain.' 


BIRDS— SUMMARY.  457 

from  this  through  natural  selection,  resulting  from  the  incon- 
venience or  danger  thus  caused. 

I  have  endeavored  in  this  chapter  to  show  that  the  arguments 
are  not  trustworthy  in  favor  of  the  view  that  weapons,  bright 
colors,  and  various  ornaments,  are  now  confined  to  the  males 
owing  to  the  conversion,  by  natural  selection,  of  the  equal  trans- 
mission of  characters  to  both  sexes,  into  transmission  to  the  male 
sex  alone.  It  is  also  doubtful  whether  tke  colors  of  many  female 
birds  are  due  to  the  preservation,  for  the  sake  of  protection  of 
variations  which  were  from  the  first  limited  in  their  transmission 
to  the  female  sex.  But  it  will  be  convenient  to  defer  any  further 
discussion  on  this  subject  until  I  treat,  in  the  following  chapter,  of 
the  differences  in  plumage  between  the  young  and  old. 


458  THE  DESCENT  OF  MAN. 


CHAPTER   XVI. 

BIRDS— concluded. 

The  immature  plumage  in  relation  to  the  character  of  the  plumage  in 
both  sexes  when  adult— Six  classes  of  cases— Sexual  differences 
between  the  males  of  closely-allied  or  representative  species — The 
female  assuming  the  characters  of  the  male — Plumage  of  the  young 
in  relation  to  the  summer  and  winter  plumage  of  the  adults— On 
the  increase  of  beauty  in  the  birds  of  the  world — Protective  color- 
ing—Conspicuously-colored  birds — Novelty  appreciated — Summary 
of  the  four  chapters  on  Birds. 

We  must  now  consider  the  transmission  of  characters,  as  limited 
by  age,  in  reference  to  sexual  selection.  The  truth  and  impor- 
tance of  the  principle  of  inheritance  at  corresponding  ages  need 
not  here  be  discussed,  as  enough  has  already  been  said  on  the  sub- 
ject. Before  giving  the  several  rather  complex  rules  or  classes  of 
cases,  under  which  the  differences  in  plumage  between  the  young 
and  the  old,  as  far  as  known  to  me,  may  be  included,  it  will  be 
well  to  make  a  few  preliminary  remarks. 

With  animals  of  all  kinds  when  the  adults  differ  in  color  from 
the  young,  and  the  colors  of  the  latter  are  not,  as  far  as  we  can 
see,  of  any  special  service,  they  may  generally  be  attributed,  like 
various  embryological  structures,  to  the  retention  of  a  former  char- 
acter. But  this  view  can  be  maintained  with  confidence,  only  when 
the  young  of  several  species  resemble  each  other  closely,  and  like- 
wise resemble  other  adult  species  belonging  to  the  same  group; 
for  the  latter  are  the  living  proofs  that  such  a  state  of  things 
was  formerly  possible.  Young  lions  and  pumas  are  marked  with 
feeble  stripes  or  rows  of  spots,  and  as  many  allied  species  both 
young  and  old  are  similarly  marked,  no  believer  in  evolution  will 
doubt  that  the  progenitor  of  the  lion  and  puma  was  a  striped  ani- 
mal, and  that  the  young  have  retained  vestiges  of  the  stripes,  like 
the  kittens  of  black  cats,  which  are  not  in  the  least  striped  when 
grown  up.  Many  species  of  deer,  which  when  mature  are  not 
spotted,  are  whilst  young  covered  with  white  spots,  as  are  like- 
wise some  few  species  in  the  adult  state.  So  again  the  young  in 
the  whole  family  of  pigs  (Suidse),  and  in  certain  rather  distantly 


BIRDS— INHERITANCE  LIMITED  BY  AGE.  459 

allied  animals,  siicli  as  the  tapir,  are  marked  with  dark  longitu- 
dinal stripes;  but  here  we  have  a  character  apparently  derived 
from  an  extinct  progenitor,  and  now  preserved  by  the  young  alone. 
In  all  such  cases  the  old  have  had  their  colors  changed  in  the 
course  of  time,  whilst  the  young  have  remained  but  little  al- 
tered, and  this  has  been  effected  through  the  principle  of  inherit- 
ance at  corresponding  ages. 

This  same  principle  applies  to  many  birds  belonging  to  various 
groups,  in  which  the  young  closely  resemble  each  other,  and  differ 
much  from  their  respective  adult  parents.  The  young  of  almost 
all  the  Gallinaceae,  and  of  some  distantly  allied  birds  such  as 
ostriches,  are  covered  with  longitudinally  striped  down;  but  this 
character  points  back  to  a  state  of  things  so  remote  that  it  hardly 
concerns  us.  Young  cross-bills  (Loxia)  have  at  first  straight 
beaks  like  those  of  other  finches,  and  in  their  immature  striated 
plumage  they  resemble  the  mature  redpole  and  female  siskin,  as 
well  as  the  young  of  the  goldfinch,  greenfinch,  and  some  other 
allied  species.  The  young  of  many  kinds  of  buntings  (Emberiza) 
resemble  one  another,  and  likewise  the  adult  state  of  the  common 
bunting,  E.  miliaria.  In  almost  the  whole  large  group  of  thrushes 
the  young  have  their  breasts  spotted — a  character  which  is  re- 
tained throughout  life  by  many  species,  but  is  quite  lost  by  others, 
as  by  the  Turdus  migratorius.  So  again  with  many  thrushes,  the 
feathers  on  the  back  are  mottled  before  they  are  moulted  for  the 
first  time,  and  this  character  is  retained  for  life  by  certain  eastern 
species.  The  young  of  many  species  of  shrikes  (Lanius),  of  some 
woodpeckers,  and  of  an  Indian  pigeon  (Chalcophaps  indicus),  are 
transversely  striped  on  the  under  surface;  and  certain  allied  spe- 
cies or  whole  genera  are  similarly  marked  when  adult.  In  some 
closely-allied  and  resplendent  Indian  cuckoos  (Chrysococcyx),  the 
mature  species  differ  considerably  from  one  another  in  color,  but 
the  young  cannot  be  distinguished.  The  young  of  an  Indian 
goose  (Sarkidiornis  melanonotus)  closely  resemble  in  plumage  an 
allied  genus,  Dendrocygna,  when  mature.^  Similar  facts  will  here- 
after be  given  in  regard  to  certain  herons.  Young  black  grouse 
(Tetrao  tetrix)  resemble  the  young  as  well  as  the  old  of  certain 
other  species,  for  instance  the  red  grouse  or  T.  scoticus.  Finally, 
as  Mr.  Blyth,  who  has  attended  closely  to  this  subject,  has  well 
remarked,  the  natural  affinities  of  many  species  are  best  exhibited 

1  In  regard  to  thrushes,  shrikes,  and  woodpeckers,  see  Mr.  Blyth,  in 
Charlesworth's  'Mag-,  of  Nat.  Hist.'  vol.  i.  1837,  p.  304;  also  footnote 
to  his  translation  of  Cuvier's  'Regne  Animal,'  p.  159.  I  give  the  case 
of  Loxia  on  Mr.  Blyth' s  information.  On  thrushes,  see,  also,  Audu- 
bon, 'Ornith.  Biography,'  vol.  ii.  p.  195.  On  Chrysococcyx  and  Chal- 
cophaps, Blyth,  as  quoted  in  Jerdon's  'Birds  of  India,'  vol.  iii.  p.  485. 
On  Sarkidiornis,  Blyth,  in  'Ibis,'  1867,  p.  175. 


460  THE  DESCENT  OF  MAN, 

in  their  immature  plumage;  and  as  the  true  affinities  of  all  organic 
beings  depend  on  their  descent  from  a  common  progenitor,  this  re- 
mark strongly  confirms  the  belief  that  the  immature  plumage 
approximately  shows  us  the  former  or  ancestral  condition  of  the 
species. 

Although  many  young  birds,  belonging  to  various  families, 
thus  give  us  a  glimpse  of  the  plumage  of  their  remote  progenitors, 
yet  there  are  many  other  birds,  both  dull-colored  and  bright-col- 
ored, in  which  the  young  closely  resemble  their  parents.  In  such 
cases  the  young  of  the  different  species  cannot  resemble  each  other 
more  closely  than  do  the  parents;  nor  can  they  strikingly  resemble 
allied  forms  when  adult.  They  give  us  but  little  insight  into  the 
plumage  of  their  progenitors,  excepting  in  so  far  that,  when  the 
young  and  the  old  are  colored  in  the  same  general  manner  through- 
out a  whole  group  of  species,  it  is  probable  that  their  progenitors 
were  similarly  colored. 

We  may  now  consider  the  classes  of  cases,  under  which  the  dif- 
ferences and  resemblances  between  the  plumage  of  the  young  and 
the  old,  in  both  sexes  or  in  one  sex  alone,  may  be  grouped.  Rules 
of  this  kind  were  first  enounced  by  Cuvier;  but  with  the  progress 
of  knowledge  they  require  some  modification  and  amplification. 
This  I  have  attempted  to  do,  as  far  as  the  extreme  complexity  of 
the  subject  permits,  from  information  derived  from  various  sour- 
ces; but  a  full  essay  on  this  subject  by  some  competent  ornith- 
ologist is  much  needed.  In  order  to  ascertain  to  what  extent 
each  rule  prevails,  I  have  tabulated  the  facts  given  in  four  great 
works,  namely,  by  Macgillivray  on  the  birds  of  Britain,  Audubon, 
on  those  of  North  America,  Jerdon  on  those  of  India,  and  Gould 
on  those  of  Australia.  I  may  here  premise,  first,  that  the  several 
cases  or  rules  graduate  into  each  other;  and,  secondly,  that  when 
the  young  are  said  to  resemble  their  parents,  it  is  not  meant  that 
they  are  identically  alike,  for  their  colors  are  almost  always  less 
vivid,  and  the  feathers  are  softer  and  often  a  different  shape. 

RULES  OR  CLASSES  OF  CASES. 

I.  "When  the  adult  male  is  more  beautiful  or  conspicuous  than 
the  adult  female,  the  young  of  both  sexes  in  their  first  plumage 
closely  resemble  the  adult  female,  as  with  the  common  fowl  and 
peacock;  or,  as  occasionally  occurs,  they  resemble  her  much  more 
closely  than  they  do  the  adult  male. 

II.  When  the  adult  female  is  more  conspicuous  than  the  adult 
male,  as  sometimes  though  rarely  occurs,  the  young  of  both  sexes 
in  their  first  plumage  resemble  the  adult  male. 

III.  When  the  adult  male  resembles  the  adult  female,  the  young 
of  both  sexes  have  a  peculiar  first  plumage  of  their  own,  as  with 
the  robin. 


BIRDS-YOUNG  LIKE  ADULT  FEMALES.  461 

IV.  When  the  adult  male  resembles  the  adult  female  the  young 
of  both  sexes  in  their  first  plumage  resemble  the  adults,  as  with 
the  kingfisher,  many  parrots,  crows,  hedge-warblers. 

V.  When  the  adults  of  both  sexes  have  a  distinct  winter  and 
summer  plumage,  whether  or  not  the  male  differs  from  the  female, 
the  young  resemble  the  adults  of  both  sexes  in  their  winter  dress, 
or  much  more  rarely  in  their  summer  dress,  or  they  resemble  the 
females  alone.  Or  the  young  may  have  an  intermediate  char- 
acter; or  again  they  may  differ  greatly  from  the  adults  in  both 
their  seasonal  plumages. 

VI.  In  some  few  cases  the  young  in  their  first  plumage  differ 
from  each  other  according  to  sex;  the  young  males  resembling 
more  or  less  closely  the  adult  males,  and  the  young  females  more 
or  less  closely  the  adult  females. 

Class  I. — In  this  class,  the  young  of  both  sexes  more  or  less 
closely  resemble  the  adult  female,  whilst  the  adult  male  differs 
from  the  adult  female,  often  in  the  most  conspicuous  manner.  In- 
numerable instances  in  all  Orders  could  be  given;  it  will  suffice  to 
call  to  mind  the  common  pheasant,  duck,  and  house-sparrow.  The 
cases  under  this  class  graduate  into  others.  Thus  the  two  sexes 
when  adult  may  differ  so  slightly,  and  the  young  so  slightly  from 
the  adults,  that  it  is  doubtful  whether  such  cases  ought  to  come 
under  the  present,  or  under  the  third  or  fourth  classes.  So  again 
the  young  of  the  two  sexes,  instead  of  being  quite  alike,  may  differ 
in  a  slight  degree  from  each  other,  as  in  our  sixth  class.  These 
transitional  cases,  however,  are  few,  or  at  least  are  not  strongly 
pronounced,  in  comparison  with  those  which  come  strictly  under 
the  present  class. 

The  force  of  the  present  law  is  well  shown  in  those  groups,  in 
which,  as  a  general  rule,  the  two  sexes  and  the  young  are  all  alike; 
for  when  in  these  groups  the  male  does  differ  from  the  female,  as 
with  certain  parrots,  kingfishers,  pigeons,  &c.,  the  young  of  both 
sexes  resemble  the  adult  female.-  We  see  the  same  fact  exhibited 
still  more  clearly  in  certain  anomalous  cases;  thus  the  male  of 

-  See,  for  instance,  Mr.  Gould's  account  ('Handbook  to  the  Birds  of 
Australia,'  vol.  i.  p.  133)  of  Cyanalcyon  (one  of  the  Kingfishers)  in 
which,  however,  the  young-  male,  though  resembling  the  adult  female, 
is  less  brilliantly  colored.  In  some  species  of  Dacelo  the  males  have 
blue  tails,  and  the  females  brown  ones;  and  Mr.  R.  B.  Sharpe  informs 
me  that  the  tail  of  the  young  male  of  D.  gaudichaudi  is  at  first  brown. 
Mr.  Gould  has  described  (ibid.  vol.  ii.  pp.  14,  20,  37)  the  sexes  and  the 
young  of  certain  black  Cockatoos  and  of  the  King  Lory,  with  which 
the  same  rule  prevails.  Also  Jerdon  ('Birds  of  India,'  vol.  i.  p.  260) 
on  the  Palaeornis  rosa,  in  which  the  young  are  more  like  th©  female 
than  the  male.  See  Audubon  ('Ornith.  Biograph.'  vol.  ii.  p.  475)  on 
the  two  sexes  and  the  young  of  Columba  passerina. 


462  THE  DESCENT  OF  MAN. 

Ileliothrix  auriculata  (one  of  the  humming-birds)  differs  con- 
spicuously from  the  female  in  having  a  splendid  gorget  and  fine 
ear-tufts,  but  the  female  is  remarkable  from  having  a  much  longer 
tail  than  that  of  the  male;  now  the  young  of  both  sexes  resemble 
(with  the  exception  of  the  breast  being  spotted  with  bronze)  the 
adult  female  in  all  other  respects,  including  the  length  of  her  tail, 
so  that  the  tail  of  the  male  actually  becomes  shorter  as  he  reaches 
maturity,  which  is  a  most  unusual  circumstance.^  Again,  the 
plumage  of  the  male  goosander  (Mergus  merganser)  is  more  con- 
spicuously colored  than  that  of  the  female,  with  the  scapular  and 
secondary  wing-feathers  much  longer;  but  differently  from  what 
occurs,  as  far  as  I  know,  in  any  other  bird,  the  crest  of  the  adult 
male,  though  broader  than  that  of  the  female,  is  considerably 
shorter,  being  only  a  little  above  an  inch  in  length;  the  crest  of 
the  female  being  two  and  a  half  inches  long.  Now  the  young  of 
both  sexes  entirely  resemble  the  adult  female,  so  that  their  crests 
are  actually  of  greater  length,  though  narrower,  than  in  the  adult 
male.* 

When  the  young  and  the  females  closely  resemble  each  other  and 
both  differ  from  the  males,  the  most  obvious  conclusion  is  that 
the  males  alone  have  been  modified.  Even  in  the  anomalous  cases 
of  the  Heliothrix  and  Mergus,  it  is  probable  that  originally  both 
adult  sexes  were  furnished — the  one  species  with  a  much  elongated 
tail,  and  the  other  v/ith  a  much  elongated  crest — these  characters 
having  since  been  partially  lost  by  the  adult  males  from  some  un- 
explained cause,  and  transmitted  in  their  diminished  state  to  their 
male  offspring  alone,  when  arrived  at  the  corresponding  age  of 
maturity.  The  belief  that  in  the  present  class  the  male  alone  has 
been  modified,  as  far  as  the  differences  between  the  male  and  the 
female  together  with  her  young  are  concerned,  is  strongly  sup- 
ported by  some  remarkable  facts  recorded  by  Mr.  Blyth,^  with  re- 
spect to  closely-allied  species  which  represent  each  other  in  dis- 
tinct countries.  For  with  several  of  these  representative  species 
the  adult  males  have  undergone  a  certain  amount  of  change  and 
can  be  distinguished;  the  females  and  the  young  from  the  distinct 
countries  being  indistinguishable,  and  therefore  absolutely  un- 
changed. This  is  the  case  with  certain  Indian  chats  (Thamno- 
bia),  with  certain  honey-suckers   (Nectarinia),  shrikes   (Tephro- 

^  I  owe  this  information  to  Mr.  Gould,  who  showed  me  the  speci- 
mens;   see,  also,  his  'Introduction  to  the  Trochilidae,'  1861,  p.  120. 

*  Macg-illivray,  'Hist.  Brit.  Birds,'  vol.  v.  pp.  207-214. 

s  See  his  admirable  paper  in  the  'Journal  of  the  Asiatic  Soc.  of  Ben- 
gal,' vol.  xix.  1850,  p.  223;  see,  also,  Jerdon,  'Birds  of  India,'  vol.  i. 
introduction,  p.  xxix.  In  regard  to  Tanysiptera,  Prof.  Schlegel  told 
Mr.  Blyth  that  he  could  distinguish  several  distinct  races,  solely  by 
comparing  the  adult  males. 


BIRDS— YOUNG  LIKE  ADULT  FEMALES.  463 

dornis),  certain  kingfishers  (Tanysiptera),  Kalij  pheasants  (Gallo- 
phasis),  and  tree-partridges   (Arboricola). 

In  some  analogous  cases,  namely  with  birds  having  a  different 
summer  and  winter  plumage,  but  with  the  tY/o  sexes  nearly  alike, 
certain  closely-allied  species  can  easily  be  distinguished  in  their 
summer  or  nuptial  plumage,  yet  are  indistinguishable  in  their 
winter  as  well  as  in  their  immature  plumage.  This  is  the  case 
with  some  of  the  closely-allied  Indian  wag-tails  or  Motacillae.  Mr. 
Swinhoe^  informs  me  that  three  species  of  Ardeola,  a  genus  of 
herons,  which  represent  one  another  on  separate  continents,  are 
"most  strikingly  different"  when  ornamented  with  their  summer 
plumes,  but  are  hardly,  if  at  all,  distinguishable  during  the  winter. 
The  young  also  of  these  three  species  in  their  immature  plumage 
closely  resemble  the  adults  in  their  winter  dress.  This  case  is  all 
the  more  interesting,  because  with  two  other  species  of  Ardeola 
both  sexes  retain,  during  the  winter  and  summer,  nearly  the  same 
plumage  as  that  possessed  by  the  three  first  species  during  the 
winter  and  in  their  immature  state;  and  this  plumage,  which  is 
common  to  several  distinct  species  at  different  ages  and  seasons, 
probably  shows  us  how  the  progenitors  of  the  genus  were  colored. 
In  all  these  cases,  the  nuptial  plumage  which  we  may  assume  was 
originally  acquired  by  the  adult  males  during  the  breeding-season, 
and  transmitted  to  the  adults  of  both  sexes  at  the  corresponding 
season,  has  been  modified,  whilst  the  winter  and  Immature  plum- 
ages have  been  left  unchanged. 

The  question  naturally  arises,  how  is  it  that  in  these  latter  cases 
the  winter  plumage  of  both  sexes,  and  in  the  former  cases  the 
plumage  of  the  adult  females,  as  well  as  the  immature  plumage 
of  the  young,  have  not  been  at  all  affected?  The  species  which 
represent  each  other  in  distinct  countries  will  almost  always  have 
been  exposed  to  somewhat  different  conditions,  but  we  can  hardly 
attribute  to  this  action  the  modification  of  the  plumage  in  the 
males  alone,  seeing  that  the  females  and  the  young,  though 
similarly  exposed,  have  not  been  affected.  Hardly  any  fact  shows 
us  more  clearly  how  subordinate  in  importance  is  the  direct  action 
of  the  conditions  of  liffe,  in  comparison  with  the  accumulation 
through  selection  of  indefinite  variations,  than  the  surprising  dif- 
ference between  the  sexes  of  many  birds;  for  both  will  have  con- 
sumed the  same  food,  and  have  been  exposed  to  the  same  climate. 
Nevertheless  we  are  not  precluded  from  believing  that  in  the 
course  of  time  new  conditions  may  produce  some  direct  effect 
either  on  both  sexes  or  from  their  constitutional  differences  chiefly 
on  one  sex.    We  see  only  that  this  is  subordinate  in  importance  to 


6  See,  also,  Mr.  Swinhoe,  in  'Ibis,'  July,  1863,  p.  131;  and  a  previous 
paper,  with  an  extract  from  a  note  by  Mr.  Blyth,  in  'Ibis,'  Jan.  1861, 
p.  25. 


464  THE  DESCENT  OF  MAIST. 

the  accumulated  results  of  selection.  Judging,  however,  from  a 
widespread  analogy,  when  a  species  migrates  into  a  new  country 
(and  this  must  precede  the  formation  of  representative  species), 
the  changed  conditions  to  which  they  will  almost  always  have 
been  exposed  will  cause  them  to  undergo  a  certain  amount  of 
fluctuating  variability.  In  this  case  sexual  selection,  which  de- 
pends on  an  element  liable  to  change — the  taste  or  admiration  of 
the  female — will  have  had  new  shades  of  color  or  other  differences 
to  act  on  and  accumulate;  and  as  sexual  selection  is  always  at 
work,  it  would  (from  what  we  know  of  the  results  on  domestic 
animals  of  man's  unintentional  selection),  be  surprising  if  animals 
inhabiting  separate  districts,  which  can  never  cross  and  thus 
blend  their  newly-acquired  characters,  were  not,  after  a  sufficient 
lapse  of  time,  differently  modified.  These  remarks  likewise  apply 
to  the  nuptial  or  summer  plumage,  whether  confined  to  the  males 
or  common  to  both  sexes. 

Although  the  females  of  the  above  closely-allied  or  representa- 
tive species,  together  with  their  young,  differ  hardly  at  all  from 
one  another,  so  that  the  males  alone  can  be  distinguished,  yet  the 
females  of  most  species  within  the  same  genus  obviously  differ 
from  each  other.  The  differences,  however,  are  rarely  as  great  as 
between  the  males.  We  see  this  clearly  in  the  whole  family  of  the 
Gallinacese:  the  females,  for  instance,  of  the  common  and  Japan 
pheasant,  and  especially  of  the  Gold  and  Amherst  pheasant — of  the 
silver  pheasant  and  the  wild  fowl — resemble  one  another  very 
closely  in  color,  whilst  the  males  differ  to  an  extraordinary  de- 
gree. So  it  is  with  the  females  of  most  of  the  Cotingidss,  Frin- 
gillidae,  and  many  other  families.  There  can  indeed  be  no  doubt 
that,  as  a  general  rule,  the  females  have  been  less  modified  than 
the  males.  Some  few  birds,  however,  offer  a  singular  and  inex- 
plicable exception;  thus  the  females  of  Paradisea  apoda  and  P. 
papuana  differ  from  each  other  more  than  do  their  respective 
males  ;^  the  female  of  the  latter  species  having  the  under  surface 
pure  white,  whilst  the  female  P.  apoda  is  deep  brown  beneath.  So, 
again,  as  I  hear  from  Professor  Newton,  the  males  of  two  species 
of  Oxynotus  (shrikes),  which  represent  each  other  in  the  Islands 
of  Mauritius  and  Bourbon,^  differ  but  little  in  color,  whilst  the 
females  differ  much.  In  the  Bourbon  species  the  female  appears 
to  have  partially  retained  an  immature  condition  of  plumage,  for 
at  first  sight  she  "might  be  taken  for  the  young  of  the  Mauritian 
"species."  These  differences  may  be  compared  with  those  in- 
explicable ones,  which  occur  independently  of  man's  selection  in 

'Wallace,   'The  Malay  Archlpelag-o,'  vol.  ii.  1869,  p.  394. 
*  These  species  are  described,  with  colored  figures,  by  M.  F.  Pollen, 
in  'Ibis,'  1866,  p.  275. 


BIRDS— YOUNG  LIKE  ADULT  FEMALES.  465 

certain  sub-breeds  of  the  game-fowl,  in  which  the  females  are  very 
difHerent,  whilst  the  males  can  hardly  be  distinguished.* 

As  I  account  so  largely  by  sexual  selection  for  the  differences 
between  the  males  of  allied  species,  how  can  the  differences  be- 
tween the  females  be  accounted  for  in  all  ordinary  cases?  We 
need  not  here  consider  the  species  which  belong  to  distinct  genera; 
for  with  these,  adaptation  to  different  habits  of  life,  and  other 
agencies,  will  have  come  into  play.  In  regard  to  the  differences 
between  the  females  within  the  same  genus,  it  appears  to  me 
almost  certain,  after  looking  through  various  large  groups,  that 
the  chief  agent  has  been  the  greater  or  less  transference  to  the 
female  of  the  characters  acquired  by  the  males  through  sexual  se- 
lection. In  the  several  British  finches,  the  two  sexes  differ  either 
very  slightly  or  considerably;  and  if  we  compare  the  females  of 
the  greenfinch,  chafiinch,  goldfinch,  bullfincn,  crossbill,  sparrow, 
&c.,  we  shall  see  that  they  differ  from  one  another  chiefly  in  the 
points  in  which  they  partially  resemble  their  respective  males; 
and  the  colors  of  the  males  may  safely  be  attributed  to  sexual  se- 
lection. With  many  gallinaceous  species  the  sexes  differ  to  an  ex- 
treme degree,  as  with  the  peacock,  pheasant,  and  fowl,  whilst  with 
other  species  there  has  been  a  partial  or  even  complete  transfer- 
ence of  character  from  the  male  to  the  female.  The  females  of 
the  several  species  of  Polyplectron  exhibit  in  a  dim  condition,  and 
chiefly  on  the  tail,  the  splendid  ocelli  of  their  males.  The  female 
partridge  differs  from  the  male  only  in  the  red  mark  on  her  breast 
being  smaller;  and  the  female  wild  turkey  only  in  her  colors  being 
much  duller.  In  the  guinea-fowl  the  two  sexes  are  indistinguish- 
able. There  is  no  improbability  in  the  plain,  though  peculiarly 
spotted  plumage  of  this  latter  bird  having  been  acquired  through 
sexual  selection  by  the  males,  and  then  transmitted  to  both  sexes; 
for  it  is  not  essentially  different  from  the  much  more  beautifully 
spotted  plumage,  characteristic  of  the  males  alone  of  the  Trago- 
pan  pheasants. 

It  should  be  observed  that,  in  some  instances,  the  transference 
of  characters  from  the  male  to  the  female  has  been  effected  ap- 
parently at  a  remote  period,  the  male  having  subsequently  under- 
gone great  changes,  without  transferring  to  the  female  any  of  his 
later-gained  characters.  For  instance,  the  female  and  the  young 
of  the  black-grouse  (Tetrao  tetrix)  resemble  pretty  closely  both 
sexes  and  the  young  of  the  red-grouse  (T.  scoticus) ;  and  we  may 
consequently  infer  that  the  black-grouse  is  descended  from  some 
ancient  species,  of  which  both  sexes  were  colored  in  nearly  the 
same  manner  as  the  red-grouse.  As  both  sexes  of  this  latter 
species  are  more  distinctly  barred  during  the  breeding  season  than 
at  any  other  time,  and  as  the  male  differs  slightly  from  the  female 

*  'Variation  of  Animals,  &c.,  under  Domestication,'  vol.  i.  p.  251. 
31 


466  THE  DESCENT  OF  MAN. 

in  his  more  strongly-pronounced  red  and  brown  tints,^"  we  may- 
conclude  that  his  plumage  has  been  influenced  by  sexual  selection, 
at  least  to  a  certain  extent.  If  so,  we  may  further  infer  that  the 
nearly  similar  plumage  of  the  female  black-grouse  was  similarly 
produced  at  some  former  period.  But  since  this  period  the  male 
black-grouse  has  acquired  his  fine  black  plumage,  with  his  forked 
and  outwardly-curled  tail-feathers;  but  of  these  characters  there 
has  hardly  been  any  transference  to  the  female  excepting  that  she 
shows  in  her  tail  a  trace  of  the  curved  fork. 

We  may  therefore  conclude  that  the  females  of  distinct  though 
allied  species  have  often  had  their  plumage  rendered  more  or  less 
different  by  the  transference  in  various  degrees,  of  characters,  ac- 
quired by  the  males  through  sexual  selection,  both  during  former 
and  recent  times.  But  it  deserves  especial  attention  that  brilliant 
colors  have  been  transferred  much  more  rarely  than  other  tints. 
For  instance,  the  male  of  the  red-throated  blue-breast  (Cyanecula 
suecica)  has  a  rich  blue  breast,  including  a  sub-triangular  red 
mark;  now  marks  of  nearly  the  same  shape  have  been  transferred 
to  the  female,  but  the  central  space  is  fulvous  instead  of  red,  and 
is  surrounded  by  mottled  instead  of  blue  feathers.  The  Gal- 
linacese  offer  many  analogous  cases;  for  none  of  the  species, 
such  as  partridges,  quails,  guinea-fowls,  &c,,  in  which  the  colors 
of  the  plumage  have  been  largely  transferred  from  the  male  to  the 
female,  are  brilliantly  colored.  This  is  well  exemplified  with  the 
pheasants,  in  which  the  male  is  generally  so  much  more  brilliant 
than  the  female;  but  with  the  Eared  and  Cheer  pheasants  (Cros- 
soptilon  auritum  and  Phasianus  wallichii)  the  sexes  closely  re- 
semble each  other  and  their  colors  are  dull.  We  may  go  so  far  as 
to  believe  that  if  any  part  of  the  plumage  in  the  males  of  these 
two  pheasants  had  been  brilliantly  colored,  it  would  not  have  been 
transferred  to  the  females.  These  facts  strongly  support  Mr.  Wal- 
lace's views  that  with  birds  which  are  exposed  to  much  danger 
during  incubation,  the  transference  of  bright  colors  from  the  male 
to  the  female  has  been  checked  through  natural  selection.  We 
must  not,  however,  forget  that  another  explanation,  before  given, 
is  possible;  namely,  that  the  males  which  varied  and  became 
bright,  whilst  they  were  young  and  inexperienced,  would  have 
been  exposed  to  much  danger,  and  would  generally  have  been  de- 
stroyed; the  older  and  more  cautious  males,  on  the  other  hand, 
if  they  varied  in  a  like  manner,  would  not  only  have  been  able  to 
survive,  but  would  have  been  favored  in  their  rivalry  with  other 
males.  Now  variations  occurring  late  in  life  tend  to  be  trans- 
mitted exclusively  to  the  same  sex,  so  that  in  this  case  extremely 
bright  tints  would  not  have  been  transmitted  to  the  females.  On 
the  other  hand,  ornaments  of  a  less  conspicuous  kind,  such  as 


i*>  Macgillivray,   'Hist.   British  Birds,'  vol.  i.   pp.  172-174. 


BIRDS- YOUNG  LIKE  ADULT  FEMALES.  467 

those  possessed  by  the  Eared  and  Cheer  pheasants,  would  not  havo 
been  dangerous,  and  if  they  appeared  during  early  youth,  would 
generally  have  been  transmitted  to  both  sexes. 

In  addition  to  the  effects  of  the  partial  transference  of  characters 
from  the  males  to  the  females,  some  of  the  differences  between  the 
females  of  closely  allied  species  may  be  attributed  to  the  direct  or 
definite  action  of  the  conditions  of  life."  With  the  males,  any 
such  action  would  generally  have  been  masked  by  the  brilliant 
colors  gained  through  sexual  selection;  but  not  so  with  the  fe- 
males. Each  of  the  endless  diversities  in  plumage,  which  we  see 
in  our  domesticated  birds  is,  of  course,  the  result  of  some  definite 
cause;  and  under  natural  and  more  uniform  conditions,  some  one 
tint,  assuming  that  it  was  in  no  way  injurious,  would  almost  cer- 
tainly sooner  or  later  prevail.  The  free  intercrossing  of  the  many 
individuals  belonging  to  the  same  species  would  ultimately  tend 
to  make  any  change  of  color,  thus  induced,  uniform  in  character. 

No  one  doubts  that  both  sexes  of  many  birds  have  had  their 
colors  adapted  for  the  sake  of  protection;  and  it  is  possible  that 
the  females  alone  of  some  species  may  have  been  modified  for  this 
end.  Although  it  would  be  a  difficult,  perhaps  an  impossible  proc- 
ess, as  shown  in  the  last  chapter,  to  convert  one  form  of  trans- 
mission into  another  through  selection,  there  would  not  be  the 
least  difficulty  in  adapting  the  colors  of  the  female,  independents 
of  those  of  the  male,  to  surrounding  objects,  through  the  accumu- 
lation of  variations  which  were  from  the  first  limited  in  their 
transmission  to  the  female  sex.  If  the  variations  were  not  thus 
limited,  the  bright  tints  of  the  male  would  be  deteriorated  or  de- 
stroyed. Whether  the  females  alone  of  many  species  have  been 
thus  specially  modified,  is  at  present  very  doubtful.  I  wish  I  could 
follow  Mr.  Wallace  to  the  full  extent;  for  the  admission  would  re- 
move some  difficulties.  Any  variations  which  were  of  no  service 
to  the  female  as  a  protection  would  be  at  once  obliterated,  instead 
of  being  lost  simply  by  not  being  selected,  or  from  free  intercross- 
ing, or  from  being  eliminated  when  transferred  to  the  male  and 
in  any  way  injurious  to  him.  Thus  the  plumage  of  the  female 
would  be  kept  constant  in  character.  It  would  also  be  a  relief  if 
we  could  admit  that  the  obscure  tints  of  both  sexes  of  many  birds 
had  been  acquired  and  preserved  for  the  sake  of  protection, — for 
example,  of  the  hedge-warbler  or  kitty-wren  (Accentor  modularis 
and  Troglodytes  vulgaris),  with  respect  to  which  we  have  no  suf- 
ficient evidence  of  the  action  of  sexual  selection.  We  ought,  how- 
ever, to  be  cautious  in  concluding  that  colors  which  appear  to  us 
dull,  are  not  attractive  to  the  females  of  certain  species;  we  should 
bear   in   mind   such   cases  as  that  of  the   common  house-spar- 

^  See,  on  this  subject,  chap,  xxiii.  in  the  'Variation  of  Animals  and 
Plants  under  Domestication.' 


468  THE  DESCENT  OF  MAN. 

row,  in  which  the  male  differs  much  from  the  female,  but  does  not 
exhibit  any  bright  tints.  No  one  probably  will  dispute  that  many 
gallinaceous  birds  which  live  on  the  open  ground,  have  acquired 
their  present  colors,  at  least  in  part,  for  the  sake  of  protection. 
"We  know  how  well  they  are  thus  concealed;  we  know  that 
ptarmigans,  whilst  changing  from  their  winter  to  their  summer 
plumage,  both  of  which  are  protective,  suffer  greatly  from  birds 
of  prey.  But  can  we  believe  that  the  very  slight  differences  in 
tints  and  markings  between,  for  instance,  the  female  black-grouse 
and  red-grouse  serve  as  a  protection?  Are  partridges,  as  they  are 
now  colored,  better  protected  than  if  they  had  resembled  quails? 
Do  the  slight  differences  between  the  females  of  the  common 
pheasant,  the  Japan  and  gold  pheasants,  serve  as  a  protection, 
or  might  not  their  plumages  have  been  interchanged  with  im- 
punity? From  what  Mr.  Wallace  has  observed  of  the  habits  of 
certain  gallinaceous  birds  in  the  East,  he  thinks  that  such  slight 
differences  are  beneficial.  For  myself,  I  will  only  say  that  I 
am  not  convinced. 

Formerly  when  I  was  inclined  to  lay  much  stress  on  protection 
as  accounting  for  the  duller  colors  of  female  birds,  it  occurred 
to  me  that  possibly  both  sexes  and  the  young  might  aboriginally 
have  been  equally  bright  colored;  but  that  subsequently,  the 
females  from  the  danger  incurred  during  incubation,  and  the 
young  from  being  inexperienced,  had  been  rendered  dull  as  a 
protection.  But  this  view  is  not  supported  by  any  evidence, 
and  is  not  probable;  for  we  thus  in  imagination  expose  during 
past  times  the  females  and  the  young  to  danger,  from  which  it 
has  subsequently  been  necessary  to  shield  their  modified  de- 
scendants. We  have,  also,  to  reduce,  through  a  gradual  process 
of  selection,  the  females  and  the  young  to  almost  exactly  the 
same  tints  and  markings,  and  to  transmit  them  to  the  correspond- 
ing sex  and  period  of  life.  On  the  supposition  that  the  females 
and  the  young  have  partaken  during  each  stage  of  the  process 
of  modification  of  a  tendency  to  be  as  brightly  colored  as  the 
males,  it  is  also  a  somewhat  strange  fact  that  the  females  have 
never  been  rendered  dull-colored  without  the  young  participating 
in  the  same  change;  for  there  are  no  instances,  as  far  as  I  can 
discover,  of  species  with  the  females  dull  and  the  young  bright 
colored.  A  partial  exception,  however,  is  offered  by  the  young 
of  certain  woodpeckers,  for  they  have  "the  whole  upper  part  of 
"the  head  tinged  with  red,"  which  afterwards  either  decreases 
into  a  mere  circular  red  line  in  the  adults  of  both  sexes,  or  quite 
disappears  in  the  adult  females.^^ 

^  Audubon,  'Ornith.  Biography,'  vol.  i.  p.  193.  Macgillivray,  'Hist. 
Brit.  Birds,'  vol.  iii.  p.  85.  See  also  the  case  before  given  of  Indoplcus 
carlotta. 


BIRDS— TOUNG  LIKE  ADULT  MALES.  469 

Finally,  with  respect  to  our  present  class  of  cases,  the  most 
probable  view  appears  to  be  that  successive  variations  in  bright- 
ness or  in  other  ornamental  characters,  occurring  in  the  males 
at  a  rather  la,te  period  of  life  have  alone  been  preserved;  and 
that  most  or  all  of  these  variations,  owing  to  the  late  period  of 
life  at  which  they  appeared,  have  been  from  the  first  transmitted 
only  to  the  adult  male  offspring.  Any  variations  in  brightness 
occurring  in  the  females  or  in  the  young,  would  have  been  of  no 
service  to  them,  and  would  not  have  been  selected;  and  more- 
over, if  dangerous,  would  have  been  eliminated.  Thus  the  fe- 
males and  the  young  will  either  have  been  left  unmodified,  or 
(as  is  much  more  common)  will  have  been  partially  modified 
by  receiving  through  transference  from  the  males  some  of  his 
successive  variations.  Both  sexes  have  perhaps  been  directly 
acted  on  by  the  conditions  of  life  to  which  they  have  long  been 
exposed:  but  the  females  from  not  being  otherwise  much  modi- 
fied, will  best  exhibit  any  such  effects.  These  changes  and  all 
others  will  have  been  kept  uniform  by  the  free  intercrossing 
of  many  individuals.  In  some  cases,  especially  with  ground 
birds,  the  females  and  the  young  may  possibly  have  been  modified, 
independently  of  the  males,  for  the  sake  of  protection,  so  as  to 
have  acquired  the  same  dull  colored  plumage. 

Class  II.  When  the  adult  female  is  more  conspicuous  than  the 
adult  male,  the  young  of  both  sexes  in  their  first  plumage  resemble 
the  adult  male. — This  class  is  exactly  the  reverse  of  the  last,  for 
the  females  are  here  brighter  colored  or  more  conspicuous  than 
the  males;  and  the  young,  as  far  as  they  are  known,  resemble 
the  adult  males  instead  of  the  adult  females.  But  the  difference 
between  the  sexes  is  never  nearly  so  great  as  with  many  birds  in 
the  first  class,  and  the  cases  are  comparatively  rare.  Mr,.  Wal- 
lace, who  first  called  attention  to  the  singular  relation  which 
exists  between  the  less  bright  colors  of  the  males  and  their 
performing  the  duties  of  incubation,  lays  great  stress  on  this 
point,^^  as  a  crucial  test  that  obscure  colors  have  been  acquired 
for  the  sake  of  protection  during  the  period  of  nesting.  A  dif- 
ferent view  seems  to  me  more  probable.  As  the  cases  are  curious 
and  not  numerous,  I  will  briefly  give  all  that  I  have  been  able  to 
find. 

In  one  section  of  the  genus  Turnix,  quail-like  birds,  the  female 
is  invariably  larger  than  the  male  (being  nearly  twice  as  large 
in  one  of  the  Australian  species),  and  this  is  an  unusual  circum- 
stance with  the  Gallinaceae.  In  most  of  the  species  the  female  is 
more  distinctly   colored   and   brighter   than  the  male,"  but  in 

13  'Westminster   Review,'   July,   1867,   and   A.    Murray,     'Journal    of 
Travel,'  1868,  p.  83. 
"  For  the  Australian  species,   see  Gould's   'Handbook,'   &c.,  vol.   it 
31 


470  THE   DESCENT  OW  MAN. 

some  few  species  the  sexes  are  alike.  In  Turnix  taigoor  of  India 
the  male  "wants  the  black  on  the  throat  and  neck,  and  the 
"whole  tone  of  the  plumage  is  lighter  and  less  pronounced  than 
"that  of  the  female."  The  female  appears  to  be  noisier,  and  is 
certainly  much  more  pugnacious  than  the  male;  so  that  the 
females  and  not  the  males  are  often  kept  by  the  natives  for 
fighting,  like  game-cocks.  As  male  birds  are  exposed  by  the 
English  bird-catchers  for  a  decoy  near  a  trap,  in  order  to  catch 
other  males  by  exciting  their  rivalry,  so  the  females  of  this 
Turnix  are  employed  in  India.  When  thus  exposed  the  females 
soon  begin  their  "loud  purring  call,  which  can  be  heard  a  long 
"way  off,  and  any  females  within  ear-shot  run  rapidly  to  the 
"spot,  and  commence  fighting  with  the  caged  bird."  In  this  way 
from  twelve  to  twenty  birds,  all  breeding  females,  may  be 
caught  in  the  course  of  a  single  day.  The  natives  assert  that 
the  females  after  laying  their  eggs  associate  in  flocks,  and  leave 
the  males  to  sit  on  them.  There  is  no  reason  to  doubt  the 
truth  of  this  assertion,  which  is  supported  by  some  observations 
made  in  China  by  Mr.  Swinhoe.^^  Mr.  Blyth  believes,  that  the 
young  of  both  sexes  resemble  the  adult  male. 

The  females  of  the  three  species  of  Painted  Snipes  (Rhynchsea, 
fig.  62)  "are  not  only  larger  but  much  more  richly  colored  than 
"the  males."^^  With  all  other  birds  in  which  the  trachea  differs 
in  structure  in  the  two  sexes  it  is  more  developed  and  complex 
in  the  male  than  in  the  female;  but  in  the  Rhynchaea  australis 
it  is  simple  in  the  male,  whilst  in  the  female  it  makes  four  dis- 
tinct convolutions  before  entering  the  lungs."  The  female  there- 
fore of  this  species  has  acquired  an  eminently  masculine  charac- 
ter. Mr.  Blyth  ascertained,  by  examining  many  specimens,  that 
the  trachea  is  not  convoluted  in  either  sex  of  R.  bengalensis, 
which  species  resembles  R.  australis  so  closely,  that  it  can  hardly 
be  distinguished  except  by  its  shorter  toes.  This  fact  is  another 
striking  instance  of  the  law  that  secondary  sexual  characters 
are  often  widely  different  in  closely-allied  forms,  though  it  is 
a  very  rare  circumstance  when  such  differences  relate  to  the 
female  sex.  The  young  of  both  sexes  of  R.  bengalensis  in  their 
first  plumage  are  said  to  resemble  the  mature  male.^^  There  is 
also   reason   to   believe  that  the   male  undertakes  the  duty  of 


pp.  178,  180,  186,  and  188.  In  the  British  Museum  specimens  of  the  Aus- 
tralian Plain-wanderer  (Pedionomus  torquatus)  may  be  seen,  showing 
similar  sexual  differences. 

1=  Jerdon,  'Birds  of  India,'  vol.  ill.  p.  596.  Mr.  Swinhoe,  in  'Ibis,' 
1865,  p.  542;  1866,  pp.  131,  405. 

^•^  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  677. 

"  Gould's  'Handbook  to  the  Birds  of  Australia,'  vol.  ii.  p.  275. 

^8  'The  Indian  Field,'  Sept.  1858,  p.  3. 


BIRDS— YOUNG  LIKE  ADULT  MALES. 


471 


incubation,  for  Mr.  Swinhoe^^  found  the  females  before  the  close 
of  the  summer  associated  in  flocks,  as  occurs  with  the  females  of 
the  Turnix. 

The  females  of  Phalaropus  fulicarius  and  P.  hj'-perboreus  are 
larger,  and  in  their  summer  plumage  "more  gayly  attired  than 


Fig.  62. 


Rhynchaea  capeiisis  (from  Brehm). 


"the  males."  But  the  difference  in  color  between  the  sexes  is 
far  from  conspicuous.  According  to  Professor  Steenstrup,  the 
male  alone  of  P.  fulicarius  undertakes  the  duty  of  incubation; 
this  is  likewise  shown  by  the  state  of  his  breast-feathers  during 
the  breeding-season.  The  female  of  the  dotterel  plover  (Eudro- 
mias  morinellus)  is  larger  than  the  male,  and  has  the  red  and 
black  tints  on  the  lower  surface,  the  white  crescent  on  the  breast. 


i»  'Ibis,'  1866,  p.  298. 


472  THE  DESCENT  OF  MAN. 

and  the  stripes  over  the  eyes,  more  strongly  pronounced.  The 
male  also  takes  at  least  a  share  in  hatching  the  eggs;  but  the 
female  likewise  attends  to  the  young.-"  I  have  not  been  able 
to  discover  whether  with  these  species  the  young  resemble  the 
adult  males  more  closely  than  the  adult  females;  for  the  com- 
parison is  somewhat  difficult  to  make  on  account  of  the  double 
moult. 

Turning  now  to  the  Ostrich  order:  the  male  of  the  common 
cassowary  (Casuarius  galeatus)  would  be  thought  by  any  one 
to  be  the  female,  from  his  smaller  size  and  from  the  appendages 
and  naked  skin  about  his  head  being  much  less  brightly  colored; 
and  I  am  informed  by  Mr.  Bartlett  that  in  the  Zoological  Gar- 
dens, it  is  certainly  the  male  alone  who  sits  on  the  eggs  and 
takes  care  of  the  young.'^  The  female  is  said  by  Mr.  T.  W. 
Wood-2  to  exhibit  during  the  breeding  season  a  most  pugnacious 
disposition;  and  her  wattles  then  become  enlarged  and  more 
brilliantly  colored.  So  again  the  female  of  one  of  the  emus 
(Dromseus  irroratus)  is  considerably  larger  than  the  male,  and 
she  possesses  a  slight  top-knot,  but  is  otherwise  indistinguishable 
in  plumage.  She  appears,  however,  "to  have  greater  power 
"when  angry  or  otherwise  excited,  of  erecting,  like  a  turkey- 
-cock, the  feathers  of  her  neck  and  breast.  She  is  usually  the 
"more  courageous  and  pugilistic.  She  makes  a  deep  hollow  gut- 
"teral  boom  especially  at  night,  sounding  like  a  small  gong.  The 
"male  has  a  slenderer  frame  and  is  more  docile,  with  no  voice 
"beyond  a  suppressed  hiss  when  angry,  or  a  croak."  He  not  only 
performs  the  whole  duty  of  incubation,  but  has  to  defend  the 
young  from  their  mother;  "for  as  soon  as  she  catches  sight  of 
"her  progeny  she  becomes  violently  agitated,  and  notwithstanding 
"the  resistance  of  the  father  appears  to  use  her  utmost  endeavors 
"to  destroy  them.  For  months  afterwards  it  is  unsafe  to  put 
"the  parents  together,  violent  quarrels  being  the  inevitable  re- 


20  For  these  several  statements,  see  Mr.  Gould's  'Birds  of  Great 
Britain.'  Prof.  Newton  informs  me  that  he  has  long-  been  convinced, 
from  his  own  observations  and  from  those  of  others,  that  the  males  of 
the  above-named  species  take  either  the  whole  or  a  large  share  of 
the  duties  of  incubation,  and  that  they  "show  much  greater  devotion 
"towards  their  young,  when  in  danger,  than  do  the  females."  So  it 
is,  as  he  informs  me,  with  Limosa  lapponica  and  some  few  other 
Waders,  in  which  the  females  are  larger  and  have  more  strongly  con- 
trasted colors  than  the  males. 

21  The  natives  of  Ceram  (Wallace,  'Malay  Archipelago,'  vol.  ii.  p.  150) 
assert  that  the  male  and  female  sit  alternately  on  the  eggs;  but  this 
assertion,  as  Mr.  Bartlett  thinks,  may  be  accounted  for  by  the  female 
visiting  the  nest  to  lay  her  eggs. 

82  -xne  Student,'  April,  1870,  p.  124. 


BIRDS— YOUNG  LIKE  ADULT  :^L\LES.  473 

*'sult,  in  whicti  the  female  generally  comes  off  conqueror."^  So 
that  with  this  emu  we  have  a  complete  reversal  not  only  of  the 
parental  and  incubating  instincts,  but  of  the  usual  moral  quali- 
ties of  the  tvro  sexes;  the  female  being  savage,  quarrelsome,  and. 
noisy,  the  males  gentle  and'good.  The  case  is  very  different  with 
the  African  ostrich,  for  the  male  is  somewhat  larger  than  the 
female  and  has  finer  plumes  with  more  strongly  contrasted  colors; 
nevertheless  he  undertakes  the  whole  duty  of  incubation.-* 

I  will  specify  the  few  other  cases  known  to  me,  in  which  the 
female  is  more  conspicuously  colored  than  the  male,  although 
nothing  is  known  about  the  manner  of  incubation.  With  the 
carrion-hawk  of  the  Falkland  Islands  (Milvago  leucurus)  I  was 
much  surprised  to  find  by  dissection  that  the  individuals,  which 
had  all  their  tints  strongly  pronounced,  with  the  cere  and  legs 
orange-colored,  were  the  adult  females;  whilst  those  with  duller 
plumage  and  gray  legs  were  the  males  or  the  young.  In  an  Aus- 
tralian tree-creeper  (Climacteris  erythrops)  the  female  differs 
from  the  male  in  "being  adorned  with  beautiful,  radiated,  rufous 
"markings  on  the  throat,  the  male  having  this  part  quite  plain." 
Lastly,  in  an  Australian  night-jar,  "the  female  always  exceeds  the 
"male  in  size  and  in  the  brilliance  of  her  tints;  the  males,  on 
"the  other  hand,  have  two  white  spots  on  the  primaries  more 
"conspicuous  than  in  the  female. -" 

We  thus  see  that  the  cases  in  which  female  birds  are  more 
conspicuously  colored  than  the  males,  with  the  young  in  their 

28  See  the  excellent  account  of  the  habits  of  this  bird  under  confine- 
ment, by  Mr.  A.  W.  Bennett,  in  'Land  and  Water,'  May,  1S6S,  p.  233. 

'■^^  Mr.  Sclater,  on  the  incubation  of  the  Struthiones,  'Proc.  Zool.  Soc.,' 
June  9,  1S63.  So  it  is  with  the  Rhea  darwinii:  Captain  Musters  says 
('At  home  with  the  Patagonians,'  1871,  p.  12S),  that  the  male  is  larg-er, 
stronger  and  swifter  than  the  female,  and  of  slig-htly  drrker  colors; 
yet  he  takes  sole  charge  of  the  eggs  and  of  the  young,  just  as  does 
the  male  of  the  common  species  of  Rhea. 

23  For  the  Milvago,  see  'Zoology  of  the  Voyage  of  the  Beagle,'  Birds, 
1S41,  p.  16.  For  the  Climacteris  and  night-jar  (Eurostopodus),  see 
Gould's  'Handbook  to  the  Birds  of  Australia,'  vol.  i.  pp.  602  and  97. 
The  New  Zealand  shieldrake  (Tadorna  variegata)  offers  a  quite  anom- 
alous case;  the  head  of  the  female  is  pure  white,  and  her  back  is 
redder  than  that  of  the  male;  the  head  of  the  male  is  of  a  rich  dark 
bronzed  color,  and  his  back  is  clothed  with  finely  penciled  slate-colored 
feathers,  so  that  altogether  he  may  be  considered  as  the  more  beau- 
tiful of  the  two.  He  is  larger  and  more  pugnacious  than  the  female, 
and  does  not  sit  on  the  eggs.  So  that  in  all  these  respects  this  species 
comes  under  our  first  class  of  cases;  but  Mr.  Sclater  ('Proc.  Zool.  Soc' 
1863,  p.  150)  was  much  surprised  to  observe  that  the  young  of  both 
sexes,  when  about  three  months  old,  resembled  in  their  dark  heads  and 
necks  the  adult  males,  instead  of  the  adult  females;  so  that  it  would 
appear  in  this  case  that  the  females  have  been  modified,  whilst  the 
males  and  the  young  have  retained  a  former  state  of  plumage. 


474  THE  DESCENT  OF  MAN. 

immature  plumage  resembling  the  adult  males  instead  of  the 
adult  females,  as  in  the  previous  class,  are  not  numerous,  though 
they  are  distributed  in  various  Orders.  The  amount  of  differ- 
ence, also,  between  the  sexes  is  incomparably  less  than  that 
which  frequently  occurs  in  the  last  class;  so  that  the  cause  of 
the  difference,  whatever  it  may  have  been,  has  here  acted  on  the 
females  either  less  energetically  or  less  persistently  than  on  the 
males  in  the  last  class.  Mr.  Wallace  believes  that  the  males 
have  had  their  colors  rendered  less  conspicuous  for  the  sake  of 
protection  during  the  period  of  incubation;  but  the  difference 
between  the  sexes  in  hardly  any  of  the  foregoing  cases  appears 
sufficiently  great  for  this  view  to  be  safely  accepted.  In  some 
of  the  cases,  the  brighter  tints  of  the  female  are  almost  confined 
to  the  lower  surface,  and  the  males,  if  thus  colored,  would  not 
have  been  exposed  to  danger  whilst  sitting  on  the  eggs.  It 
should  also  be  borne  in  mind  that  the  males  are  not  only  in  a 
slight  degree  less  conspicuously  colored  than  the  females,  but  are 
smaller  and  weaker.  They  have,  moreover,  not  only  acquired 
the  maternal  instinct  of  incubation,  but  are  less  pugnacious  and 
vociferous  than  the  females,  and  in  one  instance  have  simpler 
vocal  organs.  Thus  an  almost  complete  transposition  of  the  in- 
stincts, habits,  disposition,  color,  size,  and  of  some  points  of 
structure,  has  been  effected  between  the  two  sexes. 

Now  if  we  might  assume  that  the  males  in  the  present  class 
have  lost  some  of  that  ardor  which  is  usual  to  their  sex,  so  that 
they  no  longer  search  eagerly  for  the  females;  or,  if  we  might 
assume  that  the  females  have  become  much  more  numerous  than 
the  males — and  in  the  case  of  one  Indian  Turnix  the  females  are 
said  to  be  "much  more  commonly  met  with  than  the  males."^^ — 
then  it  is  not  improbable  that  the  females  would  have  been  led 
to  court  the  males,  instead  of  being  courted  by  them.  This  in- 
deed is  the  case  to  a  certain  extent  with  some  birds,  as  we  have 
seen  with  the  peahen,  wild  turkey,  and  certain  kinds  of  grouse. 
Taking  as  our  guide  the  habits  of  most  male  birds,  the  greater 
size  and  strength  as  well  as  the  extraordinary  pugnacity  of  the 
females  of  the  Turnix  and  emu,  must  mean  that  they  endeavor  to 
drive  away  rival  females,  in  order  to  gain  possession  of  the  male; 
and  on  this  view  all  the  facts  become  clear;  for  the  males  would 
probably  be  most  charmed  or  excited  by  the  females  which  were 
the  most  attractive  to  them  by  their  bright  colors,  other  orna- 
ments, or  vocal  powers.  Sexual  selection  would  then  do  its  work, 
steadily  adding  to  the  attractions  of  the  females;  the  males  and 
the  young  being  left  not  at  all,  or  but  little  modified. 

Class  III.  When  the  adult  male  resembles  the  adult  female, 
the  young  of  both  sexes  have  a  peculiar  first  plumage  of  their 


a«  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  598. 


BIRDS— YOUNG  LIKE  BOTH  ADULTS.  475 

own. — In  this  class  the  sexes  when  adult  resemble  each  other,  and 
differ  from  the  young.  This  occurs  with  many  birds  of  many 
kinds.  The  male  robin  can  hardly  be  distinguished  from  the 
female,  but  the  young  are  widely  different,  with  their  mottled 
dusky-olive  and  brown  plumage.  The  male  and  female  of  the 
splendid  scarlet  ibis  are  alike,  whilst  the  young  are  brown;  and 
the  scarlet  color,  though  common  to  both  sexes,  is  apparently  a 
sexual  character,  for  it  is  not  well  developed  in  either  sex  under 
confinement;  and  a  loss  of  color  often  occurs  with  brilliant 
males  when  they  are  confined.  With  many  species  of  herons 
the  young  differ  greatly  from  the  adults;  and  the  summer  plu- 
mage of  the  latter,  though  common  to  both  sexes,  clearly  has  a 
nuptial  character.  Young  swans  are  slate-colored,  whilst  the 
mature  birds  are  pure  white;  but  it  would  be  superfluous  to 
give  additional  instances.  These  differences  between  the  young 
and  the  old  apparently  depend,  as  in  the  last  two  classes,  on  the 
young  having  retained  a  former  or  ancient  state  of  plumage, 
whilst  the  old  of  both  sexes  have  acquired  a  new  one.  When  the 
adults  are  bright  colored,  we  may  conclude  from  the  remarks  just 
made  in  relation  to  the  scarlet  ibis  and  to  many  herons,  and 
from  the  analogy  of  the  species  in  the  first  class,  that  such  colors 
have  been  acquired  through  sexual  selection  by  the  nearly  ma- 
ture males;  but  that,  differently  from  what  occurs  in  the  first  two 
classes,  the  transmission,  though  limited  to  the  same  age,  has  not 
been  limited  to  the  same  sex.  Consequently,  the  sexes  when  ma- 
ture resemble  each  other  and  differ  from  the  young. 

Class  IV.  When  the  adult  male  resembles  the  adult  female, 
the  young  of  both  sexes  in  their  first  plumage  resemble  the 
adults. — In  this  class  the  young  and  the  adults  of  both  sexes, 
whether  brilliantly  or  obscurely  colored,  resemble  each  other. 
Such  cases  are,  I  think,  more  common  than  those  in  the  last  class. 
We  have  in  England  instances  in  the  kingfisher,  some  wood- 
peckers, the  jay,  magpie,  crow,  and  many  small  dull-colored  birds, 
such  as  the  hedge-warbler  or  kitty-wren.  But  the  similarity  in 
plumage  between  the  young  and  the  old  is  never  complete,  and 
graduates  away  into  dissimilarity.  Thus  the  young  of  some 
members  of  the  kingfisher  family  are  not  only  less  vividly  colored 
than  the  adults,  but  many  of  the  feathers  on  the  lower  surface 
are  edged  with  brown,-^ — a  vestige  probably  of  a  former  state  of 
the  plumage.  Frequently  in  the  same  group  of  birds,  even  within 
the  same  genus,  for  instance  in  an  Australian  genus  of  parro- 
keets  (Platycercus),  the  young  of  some  species  closely  resemble, 
whilst  the  young  of  other  species  differ  considerably,  from  their 

27  Jerdon,  'Birds  of  India,'  vol.  i.  pp.  222,  228.    Gould's  'Handbook  to 
the  Birds  of  Australia,'  vol.  i.  pp.  124,  130. 


476  THE  DESCENT  OF  MAN. 

parents  of  both  sexes,  which  are  alike.-^  Both  sexes  and  the 
young  of  the  common  jay  are  closely  similar;  but  in  the  Canada 
jay  (Perisoreus  canadensis)  the  young  differ  so  much  from  their 
parents  that  they  were  formerly  described  as  distinct  species.^'' 

I  may  remark  before  proceeding  that,  under  the  present  and 
next  two  classes  of  cases,  the  facts  are  so  complex  and  the  con- 
clusions so  doubtful,  that  any  one  who  feels  no  especial  interest 
in  the  subject  had  better  pass  them  over. 

The  brilliant  or  conspicuous  colors  which  characterize  many 
birds  in  the  present  class,  can  rarely  or  never  be  of  service  to 
them  as  a  protection;  so  that  they  have  probably  been  gained 
by  the  males  through  sexual  selection,  and  then  transferred  to 
the  females  and  the  young.  It  is,  however,  possible,  that  the 
males  may  have  selected  the  more  attractive  females;  and  if 
these  transmitted  their  characters  to  their  offspring  of  both  sexes, 
the  same  results  would  follow  as  from  the  selection  of  the  more 
attractive  males  by  the  females.  But  there  is  evidence  that  this 
contingency  has  rarely,  if  ever,  occurred  in  any  of  those  groups 
of  birds  in  which  the  sexes  are  generally  alike;  for,  if  even  a 
few  of  the  successive  variations  had  failed  to  be  transmitted  to 
both  sexes,  the  females  would  have  slightly  exceeded  the  males 
in  beauty.  Exactly  the  reverse  occurs  under  nature;  for,  in 
almost  every  large  group  in  which  the  sexes  generally  resemble 
each  other,  the  males  in  some  few  species  are  in  a  slight  degree 
more  brightly  colored  than  the  females.  It  is  again  possible  that 
tue  females  may  have  selected  the  more  beautiful  males,  these 
ma.es  having  reciprocally  selected  the  more  beautiful  females;  but 
it  is  doubtful  whether  this  double  process  of  selection  would  be 
likely  to  occur,  owing  to  the  greater  eagerness  of  one  sex  than  the 
other,  and  whether  it  would  be  more  efficient  than  selection  on  one 
side  alone.  It  is,  therefore,  the  most  probable  view  that  sexual 
selection  has  acted,  in  the  present  class,  as  far  as  ornamental 
characters  are  concerned,  in  accordance  with  the  general  rule 
throughout  the  animal  kingdom,  that  is,  on  the  males;  and  that 
these  have  transmitted  their  gradually  acquired  colors,  either 
equally  or  almost  equally,  to  their  offspring  of  both  sexes. 

Another  point  is  more  doubtful,  namely,  whether  the  suc- 
cessive variations  first  appeared  in  the  males  after  they  had  be- 
come nearly  mature,  or  whilst  quite  young.  In  either  case  sexual 
selection  must  have  acted  on  the  male  when  he  had  to  compete 
with  rivals  for  the  possession  of  the  female;  and  in  both  cases 
the  characters  thus  acquired  have  been  transmitted  to  both  sexes 
and  all   ages.    But  these  characters,   if  acquired  by  the  males 

28  Gould,  Ibid.  vol.  il.  pp.  37,  46,  56. 

2»  Audubon,  'Ornith.  Biography,'  vol.  ii.  p.  55. 


BIRDS— YOUNG   LIKE   BOTH  ADULTS.  477 

when  adult,  may  have  been  transmitted  at  first  to  the  adults 
alone,  and  at  some  subsequent  period  transferred  to  the  young. 
For  it  is  known  that,  when  the  law  of  inheritance  at  correspond- 
ing ages  fails,  the  offspring  often  inherit  characters  at  an  earlier 
age  than  that  at  which  they  first  appeared  in  their  parents.-^" 
Cases  apparently  of  this  kind  have  been  observed  with  birds  in 
a  state  of  nature.  For  instance  Mr.  Blyth  has  seen  specimens  of 
Lanius  rufus  and  of  Colymbus  glacialis  which  had  assumed  whilst 
young,  in  a  quite  anomalous  manner,  the  adult  plumage  of  their 
parents.^^  Again,  the  young  of  the  common  swan  (Cygnus  olor) 
do  not  cast  off  their  dark  feathers  and  become  white  until 
eighteen  months  or  two  years  old;  but  Dr.  F.  Forel  has  de- 
scribed the  case  of  three  vigorous  young  birds,  out  of  a  brood 
of  four,  which  were  born  pure  white.  These  young  birds  were 
not  albinoes,  as  shown  by  the  color  of  their  beaks  and  legs,  which 
nearly  resembled  the  same  parts  in  the  adults.^^ 

It  may  be  worth  while  to  illustrate  the  above  three  modes  by 
which,  in  the  present  class,  the  two  sexes  and  the  young  may 
have  come  to  resemble  each  other,  by  the  curious  case  of  the  genus 
Passer.^^  In  the  house-sparrow  (P.  domesticus)  the  male  differs 
much  from  the  female  and  from  the  young.  The  young  and  the 
females  are  alike,  and  resemble  to  a  large  extent  both  sexes  and 
the  young  of  the  sparrow  of  Palestine  (P.  br'achydactylus),  as  well 
as  some  allied  species.  We  may  therefore  assume  that  the  female 
and  young  of  the  house-sparrow  approximately  show  us  the  plu- 
mage of  the  progenitor  of  the  genus.  Now  with  the  tree-sparrow 
(P.  montanus)  both  sexes  and  the  young  closely  resemble  the 
male  of  the  house-sparrow;  so  that  they  have  all  been  modified 
in  the  same  manner,  and  all  depart  from  the  typical  coloring  of 
their  early  progenitor.  This  may  have  been  effected  by  a  male 
ancestor  of  the  tree-sparrov/  having  varied,  firstly,  when  nearly 
mature;  or  secondly,  whilst  quite  young,  and  by  having  in  either 
case  transmitted  his  modified  plumage  to  the  females  and  the 
young;  or,  thirdly,  he  may  have  varied  when  adult  and  trans- 
mitted his  plumage  to  both  adult  sexes,  and,  owing  to  the  failure 
of  the  law  of  inheritance  at  corresponding  ages,  at  some  subse- 
quent period  to  his  young. 

It  is  impossible  to  decide  which  of  these  three  modes  has  gen- 

30  'Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
p.  79. 

31  Cliarlesworth's  'Mag.  of  Nat.  Hist.'  vol.  i.  1837,  pp.  305,  306. 

32  'Bulletin  de  la  Soc.  Vaudoise  des  Sc.  Nat.'  vol.  x.  1869,  p.  132.  The 
young-  of  the  Polish  swan  Cygnus  immutabilis  of  Tarrell,  are  always 
white;  but  this  species,  as  Mr.  Sclater  informs  me,  is  believed  to  be 
nothing  more  than  a  variety  of  the  domestic  swan  (Cygnus  olor). 

32  I  am  indebted  to  Mr.  Blyth  for  information  in  regard  to  this 
genus.    The  sparrow  of  Palestine  belongs  to  the  sub-genus  Petronia. 


478  THE  DESCENT  OF  MAN. 

erally  prevailed  throughout  the  present  class  of  cases.  That  the 
males  varied  whilst  young,  and  transmitted  their  variations  to 
their  offspring  of  both  sexes,  is  the  most  probable.  I  may  here 
add  that  I  have,  with  little  success,  endeavored,  by  consulting 
various  works,  to  decide  how  far  the  period  of  variation  in  birds 
has  generally  determined  the  transmission  of  characters  to  one 
sex  or  to  both.  The  two  rules,  often  referred  to  (namely,  that 
variations  occurring  late  in  life  are  transmitted  to  one  and  the 
same  sex,  whilst  those  which  occur  early  in  life  are  transmitted 
to  both  sexes),  apparently  hold  good  in  the  first,^*  second,  and 
fourth  classes  of  cases;  but  they  fail  in  the  third,  often  in  the 
fifth,'^  and  in  the  sixth  small  class.  They  apply,  however,  as  far 
as  I  can  judge,  to  a  considerable  majority  of  the  species;  and  we 
must  not  forget  the  striking  generalization  by  Dr.  W.  Marshall 
with  respect  to  the  protuberances  on  the  heads  of  birds.  Whether 
or  not  the  two  rules  generally  hold  good,  we  may  conclude  from 
the  facts  given  in  the  eighth  chapter,  that  the  period  of  variation 
is  one  important  element  in  determining  the  form  of  transmis- 
sion. 

With  birds  it  is  difficult  to  decide  by  what  standard  we  ought 
to  judge  of  the  earliness  or  lateness  of  the  period  of  variation, 
whether  by  the  age  in  reference  to  the  duration  of  life,  or  to 
the  pov/er  of  reproduction,  or  to  the  number  of  moults  through 
which  the  species  passes.  The  moulting  of  birds,  even  within 
the  same  family,  sometimes  differs  much  without  any  assignable 
cause.  Some  birds  moult  so  early,  that  nearly  all  the  body  feath- 
ers are  cast  off  before  the  first  wing-feathers  are  fully  grown; 
and  we  cannot  believe  that  this  was  the  primordial  state  of 
things.  When  the  period  of  moulting  has  been  accelerated,  the 
age  at  which  the  colors  of  the  adult  plumage  are  first  developed 
will  falsely  appear  to  us  to  be  earlier  than  it  really  is.  This 
may  be  illustrated  by  the  practice  followed  by  some  bird-fanciers, 
who  pull  out  a  few  feathers  from  the  breast  of  nestling  bullfinches, 
and  from  the  head  or  neck  of  young  gold-pheasants,  in  order  to 
ascertain  their  sex;    for  in  the  males,  these  feathers  are  imme- 

34  For  instance,  the  males  of  Tanagra  aestiva  and  Fringilla  cyanea 
require  three  years,  the  male  of  Fringilla  ciris  four  years,  to  com- 
plete their  beautiful  plumage.  (See  Audubon,  'Ornith.  Biography,'  vol. 
i.  PP,  233,  280,  378.)  The  Harlequin  duck  takes  three  years  (ibid.  vol. 
iii.  p.  614).  The  male  of  the  Gold  pheasant,  as  I  hear  from  Mr.  Jenner 
Weir,  can  be  distinguished  from  the  female  when  about  three  months 
old,  but  he  does  not  acquire  his  full  splendor  until  the  end  of  the 
September  in  the  following  year. 

35  Thus  the  Ibis  tantalus  and  Grus  americanus  take  four  years, 
the  Flamingo  several  years,  and  the  Ardea  ludovicana  two  years, 
before  they  acquire  their  perfect  plumage.  See  Audubon,  ibid.  vol.  i. 
p.  221;  vol.  iii.  pp.  133,  139,  211. 


BIRDS— YOUNG  LIKE  BOTH  ADULTS.  479 

diately  replaced  by  colored  ones.^'*'  The  actual  duration  of  life 
is  known  in  but  few  birds,  so  tbat  we  can  hardly  judge  by  this 
standard.  And,  with  reference  to  the  period  at  which  the  power 
of  reproduction  is  gained,  it  is  a  remarkable  fact  that  various 
birds  occasionally  breed  whilst  retaining  their  immature  plum- 
age." 

The  fact  of  birds  breeding  in  their  immature  plumage  seems  op- 
posed to  the  belief  that  sexual  selection  has  played  as  important 
a  part,  as  I  believe  it  has,  in  giving  ornamental  colors,  plumes, 
&c.,  to  the  males,  and,  by  means  of  equal  transmission,  to  the 
females  of  many  species.  The  objection  would  be  a  valid  one, 
if  the  younger  and  less  ornamented  males  were  as  successful  in 
winning  females  and  propagating  their  kind,  as  the  older  and 
more  beautiful  males.  But  we  have  no  reason  to  suppose  that 
this  is  the  case.  Audubon  speaks  of  the  breeding  of  the  immature 
males  of  Ibis  tantalus  as  a  rare  event,  as  does  Mr.  Swinhoe,  in 
regard  to  the  immature  males  of  Oriolus.^"*  If  the  young  of  any 
species  in  their  immature  plumage  were  more  successful  in  win- 
ning partners  than  the  adults,  the  adult  plumage  would  probably 
soon  be  lost,  as  the  males  would  prevail,  which  retained  their 
immature  dress  for  the  longest  period,  and  thus  the  character  of 
the  species  would  ultimately  be  modified.^'    If,  on  the  other  hand, 


38  Mr.  Blyth,  in  Charlesworth's  'Mag-,  of  Nat.  Hist.'  vol.  i.  1837,  p. 
300.    Mr.  Bartlett  has  informed  me  in  regard  to  grold-pheasants. 

37  I  have  noticed  the  following-  cases  in  Audubon's  'Ornith.  Bio- 
g-raphy.'  The  redstart  of  America  (Muscapica  ruticilla,  vol.  i.  p.  203). 
The  Ibis  tantalus  takes  four  years  to  come  to  full  maturity,  but  some- 
times breeds  in  the  second  year  (vol.  iii.  p.  133).  The  Grus  ameri- 
canus  takes  the  same  time,  but  breeds  before  acquiring-  its  full 
plumage  (vol.  iii.  p.  211).  The  adults  of  Ardea  caerulea  are  blue,  and 
the  young  white;  and  white,  mottled,  and  mature  blue  birds  may  all 
be  seen  breeding  together  (vol.  iv.  p.  58):  but  Mr.  Blyth  informs  me 
that  certain  herons  apparently  are  dimorphic,  for  white  and  colored 
individuals  of  the  same  age  may  be  observed.  The  Harlequin  duck 
(Anas  histrionica,  Linn.)  takes  three  years  to  acquire  its  full  plum- 
age, though  many  bii'ds  breed  in  the  second  year  (vol.  iii.  p.  614).  The 
White-headed  eagle  (Falco  leucocephalus,  vol.  iii.  p.  210)  is  likewise 
known  to  breed  in  its  immature  state.  Some  species  of  Oriolus  (ac- 
cording to  Mr.  Blyth  and  Mr.  Swinhoe,  in  'Ibis,'  July  1863,  p.  68)  like- 
wise breed  before  they  attain  their  full  plumage. 

38  See  the  last  foot-note. 

39  Other  animals,  belonging  to  quite  distinct  classes,  are  either 
habitually  or  occasionally  capable  of  breeding  before  they  have  fully 
acquired  their  adult  characters.  This  is  the  case  with  the  young 
males  of  the  salmon.  Several  amphibians  have  been  known  to  breed 
whilst  retaining  their  larval  structure.  Fritz  Muller  has  shown 
('Facts  and  Arguments  for  Darwin,'  Eng.  trans.  1869,  p.  79)  that  the 
EPales  of  several  amphipod  crustaceans  become  sexually  mature  whilst 
>oung;  and  I  infer  that  this  is  a  case  of  premature  breeding,  because 


480  THE  DESCENT  OF  MAN. 

the  young  never  succeeded  in  obtaining  a  female,  the  habit  of 
early  reproduction  would  perhaps  be  sooner  or  later  eliminated, 
from  being  superfluous  and  entailing  waste  of  power. 

The  plumage  of  certain  birds  goes  on  increasing  in  beauty 
during  many  years  after  they  are  fully  mature;  this  is  the  case 
with  the  train  of  the  peacock,  with  some  of  the  birds  of  paradise, 
and  with  the  crest  and  plumes  of  certain  herons,  for  instance,  the 
Ardea  ludovicana.*°  But  it  is  doubtful  whether  the  continued  de- 
velopment of  such  feathers  is  the  result  of  the  selection  of  suc- 
cessive beneficial  variations  (though  this  is  the  most  probable 
view  with  birds  of  paradise)  or  merely  of  continuous  growth. 
Most  fishes  continue  increasing  in  size,  as  long  as  they  are  in 
good  health  and  have  plenty  of  food;  and  a  somewhat  similar 
law  may  prevail  with  the  plumes  of  birds. 

Class  V.  When  the  adults  of  both  sexes  have  a  distinct  winter 
and  summer  plumage,  whether  or  not  the  male  differs  from  the 
female,  the  young  resemble  the  adults  of  both  sexes  in  their 
winter  dress,  or  much  more  rarely  in  their  summer  dress,  or 
they  resemble  the  females  alone.  Or  the  young  may  have  an 
intermediate  character;  or,  again,  they  may  differ  greatly  from 
the  adults  in  both  their  seasonal  plumages. — The  cases  in  this 
class  are  singularly  complex;  nor  is  this  surprising,  as  they  de- 
pend on  inheritance,  limited  in  a  greater  or  less  degree  in  three 
different  ways,  namely,  by  sex,  age,  and  the  season  of  the  year. 
In  some  cases  the  individuals  of  the  same  species  pass  through  at 
least  five  distinct  states  of  plumage.  With  the  species,  in  which 
the  male  differs  from  the  female  during  the  summer  season  alone, 
or,  which  is  rarer,  during  both  seasons,*^  the  young  generally  re- 
semble the  females, — as  with  the  so-called  goldfinch  of  North 
America,  and  apparently  with  the  splendid  Maluri  of  Australia.** 
With  those  species,  the  sexes  of  which  are  alike  during  both 
the  summer  and  winter,  the  young  may  resemble  the  adults,  first- 
ly, in  their  winter  dress;    secondly,  and  this  is  of  much  rarer 


they  have  not  as  yet  acquired  their  fully-developed  claspers.  All 
such  facts  are  highly  interesting-,  as  bearing  on  one  means  by  which 
species  may  undergo  great  modifications  of  character. 

*o  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  507,  on  the  peacock.  Dr. 
Marshall  thinks  that  the  older  and  more  brilliant  males  of  birds  of 
paradise  have  an  advantage  over  the  younger  males;  see  'Archives 
Neerlandaises,*  torn.  vi.  1871.— On  Ardea,  Audubon,  ibid.  vol.  iii.  p.  139. 

*i  For  illustrative  cases  see  vol.  iv.  of  Macgillivray's  'Hist.  Brit. 
Birds;'  on  Tringa,  &c.,  pp.  229,  271;  on  the  Machetes,  p.  172;  on  the 
Charadrius  hiaticula,  p.  118;  on  the  Charadrius  pluvialis,  p.  94. 

*2  For  the  goldfinch  of  N.  America,  iTingilla  tristis,  Linn,,  see  Au- 
dubon. 'Ornith.  Biography,'  vol.  i.  p.  172.  For  the  Maluri,  Gould's 
•Handbook  to  the  Birds  of  Australia,'  vol.  i.  p.  318. 


BIRDS— YOUNG  LIKE  BOTH  ADULTS.  481 

occurrence,  in  their  summer  dress;  thirdly,  they  may  he  inter- 
mediate between  these  two  states;  and,  fourthly,  they  may  differ 
greatly  from  the  adults  at  all  seasons.  We  have  an  instance  of 
the  first  of  these  four  cases  in  one  of  the  egrets  of  India  (Buphus 
coromandus),  in  which  the  young  and  the  adults  of  both  sexes 
are  white  during  the  winter,  the  adults  becoming  golden-buff 
during  the  summer.  With  the  gaper  (Anastomus  oscitans)  of 
India  we  have  a  similar  case,  but  the  colors  are  reversed:  for 
the  young  and  the  adults  of  both  sexes  are  gray  and  black  during 
the  winter,  the  adults  becoming  white  during  the  summer.*^  As 
an  instance  of  the  second  case,  the  young  of  the  razor-bill  (Alca 
torda,  Linn.),  in  an  early  state  of  plumage,  are  colored  like  the 
adults  during  the  summer;  and  the  young  of  the  white-crowned 
sparrow  of  North  America  (Fringilla  leucophrys),  as  soon  as 
fledged,  have  elegant  white  stripes  on  their  heads,  which  are  lost 
by  the  young  and  the  old  during  the  winter.**  With  respect  to 
the  third  case,  namely,  that  of  the  young  having  an  intermediate 
character  between  the  summer  and  winter  adult  plumages,  Yar- 
rell*^  insists  that  this  occurs  with  many  waders.  Lastly,  in  regard 
to  the  young  differing  greatly  from  both  sexes  in  their  adult  sum- 
mer and  winter  plumages,  this  occurs  with  some  herons  and  egrets 
of  North  America  and  India, — the  young  alone  being  white. 

I  will  make  only  a  few  remarks  on  these  complicated  cases. 
When  the  young  resemble  the  females  in  their  summer  dress,  or 
the  adults  of  both  sexes  in  their  winter  dress,  the  cases  differ 
from  those  given  under  Classes  I.  and  III.  only  in  the  characters 
originally  acquired  by  the  males  during  the  breeding-season,  hav- 
ing been  limited  in  their  transmission  to  the  corresponding  sea- 
son. When  the  adults  have  a  distinct  summer  and  winter  plum- 
age, and  the  young  differ  from  both,  the  case  is  more  difficult  to 
understand.  We  may  admit  as  probable  that  the  young  have 
retained  an  ancient  state  of  plumage;  we  can  account  by  sexual 
selection  for  the  summer  or  nuptial  plumage  of  the  adults,  but 
how  are  we  to  account  for  their  distinct  winter  plumage?  If  we 
could  admit  that  this  plumage  serves  in  all  cases  as  a  protection, 
its  acquirement  would  be  a  simple  affair;  but  there  se'ems  no 
good  reason  for  this  admission.  It  may  be  suggested  that  the 
widely  different  conditions  of  life  during  the  winter  and  summer 


*?•  I  am  indebted  to  Mr.  Blyth  for  information  as  to  the  Buphus;  see 
also  Jerdon,  'Birds  of  India,'  vol.  iii.  p.  749.  On  the  Anastomus,  see 
Blyth,  in  'Ibis,'  1867,  p.  173. 

**  On  the  Alca,  see  Macg-illivray,  'Hist,  of  Brit.  Birds.'  vol.  v.  p.  347. 
On  the  Fringilla  leucophrys,  Audubon,  ibid.  vol.  ii.  p.  89.  I  shall  have 
hereafter  to  refer  to  the  young-  of  certain  herons  and  egrets  being 
white. 

^  'History  of  British  Birds,'  vol.  i.  1839,  p.  159. 
32 


482  THE  DESCENT  OP  MAN. 

have  acted  in  a  direct  manner  on  the  plumage;  this  mayhave 
had  some  effect,  but  I  have  not  much  confidence  in  so  great  a 
difference  as  we  sometimes  see  between  the  two  plumages  having 
been  thus  caused.  A  more  probable  explanation  is,  that  an  an- 
cient style  of  plumage,  partially  modified,  through  the  transfer- 
ence of  some  characters  from  the  summer  plumage,  has  been 
retained  by  the  adults  during  the  winter.  Finally,  all  the  cases 
in  our  present  class  apparently  depend  on  characters  acquired  by 
the  adult  males,  having  been  variously  limited  in  their  transmis- 
sion according  to  age,  season,  and  sex;  but  it  would  not  be  worth 
while  to  attempt  to  follow  out  these  complex  relations. 

Class  VI.  The  young  in  their  first  plumage  differ  from  each 
other  according  to  sex;  the  young  males  resembling  more  or  less 
closely  the  adult  males,  and  the  young  females  more  or  less  closely 
the  adult  females. — The  cases  in  the  present  class,  though  occur- 
ring in  various  groups,  are  not  numerous;  yet  it  seems  the  most 
natural  thing  that  the  young  should  at  first  somewhat  resemble 
the  adults  of  the  same  sex,  and  gradually  become  more  and  more 
like  them.  The  adult  male  blackcap  (Sylvia  atricapilla)  has  a 
black  head,  that  of  the  female  being  reddish-brown;  and  I  am 
informed  by  Mr.  Blyth,  that  the  young  of  both  sexes  can  be  dis- 
tinguished by  this  character  even  as  nestlings.  In  the  family  of 
thrushes  an  unusual  Jilimber  of  similar  cases  have  been  noticed; 
thus,  the  male  blackbird  (Turdus  merula)  can  be  distinguished 
in  the  nest  from  the  female.  The  two  sexes  of  the  mocking  bird 
(Turdus  polyglottus,  Linn.)  differ  very  little  from  each  other,  yet 
the  males  can  easily  be  distinguished  at  a  very  early  age  from  the 
females  by  showing  more  pure  white.^"  The  males  of  a  forest- 
thrush  and  of  a  rock-thrush  (Orocetes  erythrogastra  and  Petro- 
cincla  cyanea)  have  much  of  their  plumage  of  a  fine  blue,  whilst 
the  females  are  brown;  and  the  nestling  males  of  both  species 
have  their  main  wing  and  tail-feathers  edged  with  blue,  whilst 
those  of  the  female  are  edged  with  browm.*^  In  the  young  black- 
bird the  wing  feathers  assume  their  mature  character  and  be- 
come black  after  the  others;  on  the  other  hand,  in  the  two  spe- 
cies just  named  the  wing-feathers  become  blue  before  the  others. 
The  most  probable  view  With  reference  to  the  cases  in  the  present 
class  is  that  the  males,  differently  from  what  occurs  in  Class  I., 
have  transmitted  their  colors  to  their  male  offspring  at  an  earlier 
age  than  that  at  which  they  were  first  acquired;  for,  if  the  males 

*8  Audubon,  'Ornith.  Biography,'  vol.  i.  p.  113. 

*'  Mr.  C.  A.  Wright,  in  'Ibis,'  vol.  vi.  18S4,  p.  65.  Jerdon,  'Birds  of 
Inclia,'  vol.  i.  p.  515.  See  also  on  the  blackbird,  Blyth  in  Charles- 
worth's  'Mag.  of  Nat.  History,'  vol.  i.  1837,  p.  113. 


BIRDS— YOUNG  LIKE  BOTH  ADULTS.  483 

had  varied  whilst  quite  young,  their  characters  would  prohably 
have  been  transmitted  to  both  sexes.'*^ 

In  Aithurus  polytmus,  a  humming-bird,  the  male  is  splendidly 
colored  black  and  green,  and  two  of  the  tail-feathers  are  immense- 
ly lengthened;  the  female  has  an  ordinary  tail  and  inconspicuous 
colors;  now  the  young  males,  instead  of  resembling  the  adult 
female,  in  accordance  with  the  common  rule,  begin  from  the  first 
to  assume  the  colors  proper  to  their  sex,  and  their  tail-feathers 
soon  become  elongated.  I  owe  this  information  to  Mr.  Gould,  who 
has  given  me  the  following  more  striking  and  as  yet  unpublished 
case.  Two  humming-birds  belonging  to  the  genus  Eustephanus, 
both  beautifully  colored,  inhabit  the  small  island  of  Juan  Fernan- 
dez, and  have  always  been  ranked  as  specifically  disitinct.  But  it 
has  lately  been  ascertained  that  the  one,  which  is  of  a  rich  chest- 
nut brown  color  with  a  golden-red  head,  is  the  male,  whilst  the 
other,  which  is  elegantly  variegated  with  green  and  white  with  a 
metallic  green  head  is  the  female.  Now  the  young  from  the  first 
somewhat  resemble  the  adults  of  the  corresponding  sex,  the  re- 
semblance gradually  becoming  more  and  more  complete. 

In  considering  this  last  case,  if  as  before  we  take  the  plumage 
of  the  young  as  our  guide,  it  would  appear  that  both  sexes  have 
been  rendered  beautiful  independently;  and  not  that  one  sex 
has  partially  transferred  its  beauty  to  the  other.  The  male  ap- 
parently has  acquired  his  bright  colors  through  sexual  selection  in 
the  same  manner  as,  for  instance,  the  peacock  or  pheasant  in  our 
first  class  of  cases;  and  the  female  in  the  same  manner  as  the 
female  Rhynchsea  or  Turnix  in  our  second  class  of  cases.  But 
there  is  much  difficulty  in  understanding  how  this  could  have 
been  effected  at  the  same  time  with  the  two  sexes  of  the  same 
species.  Mr.  Salvin  states,  as  we  have  seen  in  the  eighth  chapter, 
that  with  certain  humming-birds  the  males  greatly  exceed  the 
females  in  number,  whilst  with  other  species  inhabiting  the  same 
country  the  females  greatly  exceed  the  males.  If,  then,  we  might 
assume  that  during  some  former  lengthened  period  the  males  of 
the  Juan  Fernandez  species  had  greatly  exceeded  the  females  in 
number,  but  that  during  another  lengthened  period  the  females 
had  far  exceeded  the  males,  we  could  understand  how  the  males 
at  one  time,  and  the  females  at  another,  might  have  been  ren- 


*s  The  following  additional  cases  may  be  mentioned ;  the  young  males 
of  Tanagra  rubra  can  be  distinguished  from  the  young  females  (Au- 
dubon, 'Ornith.  BiogTaphy,'  vol.  iv.  p.  392),  and  so  it  is  with  the  nest- 
lings of  a  blue  nuthatch,  Dendrophila  frontalis  of  India  (Jerdon, 
'Birds  of  India,'  vol.  i.  p.  389).  Mr.  Blyth  also  informs  me  that  the  sexes 
of  the  stanechat,  Saxicola  rubicola,  are  distinguishable  at  a  very 
early  age.  Mr.  Salvin  gives  ('Proc.  Zoolog.  Soc'  1870,  p.  206),  the  case 
of  a  humming-bird,  like  the  following  one  of  Eustephanus. 


484  THE  DESCENT  OF  MAN. 

dered  beautiful  by  the  selection  of  the  brighter-colored  individuals 
of  either  sex;  both  sexes  transmitting  their  characters  to  their 
young  at  a  rather  earlier  age  than  usual.  Whether  this  is  the 
true  explanation  I  will  not  pretend  to  say;  but  the  case  is  too 
remarkable  to  be  passed  over  without  notice. 

We  have  now  seen  in  all  six  classes,  that  an  intimate  relation 
exists  between  the  plumage  of  the  young  and  the  adults,  either 
of  one  sex  or  both.  These  relations  are  fairly  well  explained 
on  the  principle  that  one  sex — this  being  in  the  great  majority 
of  cases  the  male — first  acquired  through  variation  and  sexual 
selection  bright  colors  or  other  ornaments,  and  transmitted  them 
in  various  ways,  in  accordance  with  the  recognized  laws  of  in- 
heritance. Why  variations  have  occurred  at  different  periods  of 
life,  even  sometimes  with  species  of  the  same  group,  we  do  not 
know,  but  with  respect  to  the  form  of  transmission,  one  important 
determining  cause  seems  to  be  the  age  at  which  the  variations 
first  appear. 

From  the  principle  of  inheritance  at  corresponding  ages,  and 
from  any  variations  in  color  which  occurred  in  the  males  at  an 
early  age  not  being  then  selected — on  the  contrary  being  often 
eliminated  as  dangerous — whilst  similar  variations  occurring 
at  or  near  the  period  of  reproduction  have  been  preserved,  it 
follows  that  the  plumage  of  the  young  will  often  have  been  left 
unmodified,  or  but  little  modified.  We  thus  get  some  insight 
into  the  coloring  of  the  progenitors  of  our  existing  species.  In 
a  vast  number  of  species  in  five  out  of  our  six  classes  of  cases,  the 
adults  of  one  sex  or  of  both  are  bright  colored,  at  least  during 
the  breeding-season,  whilst  the  young  are  invariably  less  brightly 
colored  than  the  adults,  or  are  quite  dull  colored;  for  no  in- 
stance is  known,  as  far  as  I  can  discover,  of  the  young  of  dull- 
colored  species  displaying  bright  colors,  or  of  the  young  of  bright- 
colored  species  being  more  brilliant  than  their  parents.  In  the 
fourth  class,  however,  in  which  the  young  and  the  old  resemble 
each  other,  there  are  many  species  (though  by  no  means  all), 
of  which  the  young  are  bright-colored,  and  as  these  form  whole 
groups,  we  may  infer  that  their  early  progenitors  were  likewise 
bright.  With  this  exception,  if  we  look  to  the  birds  of  the  world, 
it  appears  that  their  beauty  has  been  much  increased  since  that 
period,  of  which  their  immature  plumage  gives  us  a  partial  reC' 
ord. 

On  the  Color  of  the  Plumage  in  relation  to  Protection. — It  will 
have  been  seen  that  I  cannot  follow  Mr.  Wallace  in  the  belief 
that  dull  colors,  when  confined  to  the  females,  have  been  in 
most  cases  specially  gained  for  the  sake  of  protection.  There 
can,  however,  be  no  doubt,  as  formerly  remarked,  that  both  sexes 


BIRDS-COLOR  AND  PROTECTION.  485 

of  many  birds  have  had  their  colors  modified,  so  as  to  escape 
the  notice  of  their  enemies;  or  in  some  instances,  so  as  to  ap- 
proach their  prey  unobserved,  just  as  owls  have  had  their  plum- 
age rendered  soft,  that  their  flight  may  not  be  overheard.  Mr. 
Wallace  remarks*^  that  "it  is  only  in  the  tropics,  among  forests 
"which  never  lose  their  foliage,  that  we  find  whole  groups  of  birds, 
"whose  chief  color  is  green."  It  will  be  admitted  by  every  one, 
who  has  ever  tried,  how  difficult  it  is  to  distinguish  parrots  in  a 
leaf-covered  tree.  Nevertheless,  we  must  remember  that  many 
parrots  are  ornamented  with  crimson,  blue,  and  orange  tints 
which  can  hardly  be  protective.  Woodpeckers  are  eminently  ar- 
boreal, but  besides  green  species,  there  are  many  black,  and 
black-and-white  kinds — all  the  species  being  apparently  exposed 
to  nearly  the  same  dangers.  It  is  therefore  probable  that  with 
tree-haunting  birds,  strongly-pronounced  colors  have  been  ac- 
quired through  sexual  selection,  but  that  a  green  tint  has  been 
acquired  oftener  than  any  other,  from  the  additional  advantage  of 
protection. 

In  regard  to  birds  which  live  on  the  ground,  every  one  admits 
that  they  are  colored  so  as  to  imitate  the  surrounding  surface. 
How  diflScult  it  is  to  see  a  partridge,  snipe,  woodcock,  certain 
plovers,  larks,  and  night-jars  when  crouched  on  ground.  Animals 
inhabiting  deserts  offer  the  most  striking  cases,  for  the  bare 
surface  affords  no  concealment,  and  nearly  all  the  smaller  quad- 
rupeds, reptiles,  and  birds  depend  for  safety  on  their  colors.  Mr. 
Tristram  has  remarked  in  regard  to  the  inhabitants  of  the  Sahara, 
that  all  are  protected  by  their  "isabelline  or  sand-color."^"  Call- 
ing to  my  recollection  the  desert-birds  of  South  America,  as  well 
as  most  of  the  ground-birds  of  Great  Britain,  it  appeared  to  me 
that  both  sexes  in  such  cases  are  generally  colored  nearly  alike. 
Accordingly,  I  applied  to  Mr.  Tristram  with  respect  to  the  birds 
of  the  Sahara,  and  he  has  kindly  given  me  the  following  informa- 
tion. There  are  twenty-six  species  belonging  to  fifteen  genera, 
which  manifestly  have  their  plumage  colored  in  a  protective  man- 
ner; and  this  coloring  is  all  the  more  striking,  as  with  most  of 
these  birds  it  differs  from  that  of  their  congeners.  Both  sexes 
of  thirteen  out  of  the  twenty-six  species  are  colored  in  the  same 
manner;  but  these  belong  to  genera  in  which  this  rule  commonly 
prevails,  so  that  they  tell  us  nothing  about  the  protective  colors 
being  the  same  in  both  sexes  of  desert-birds.  Of  the  other  thir- 
teen species,  three  belong  to  genera  in  which  the  sexes  usually 
differ  from  each  other,  yet  here  they  have  the  sexes  alike.    In  the 

**  'Westminster  Review,'  July,  1867,  p.  5. 

^  'Ibis,'  1S59,  vol.  i.  p.  429,  et  seq.    Dr.  Rohlfs,  however,  remarks  to 
me  in  a  letter  that,  according  to  his   experience  of  the  Sahara,   this 
statement  is  too  strong. 
32 


486  THE  DESCENT  OF  MAN. 

remaining  ten  species,  the  male  differs  from  the  female;  but  the 
difference  is  confined  chiefly  to  the  under  surface  of  the  plumage, 
-which  is  concealed  when  the  bird  crouches  on  the  ground;  the 
head  and  back  being  of  the  same  sand-colored  hue  in  the  two 
sexes.  So  that  in  these  ten  species  the  upper  surfaces  of  both 
sexes  have  been  acted  on  and  rendered  alike,  through  natural 
selection,  for  the  sake  of  protection;  whilst  the  lower  surfaces  of 
the  males  alone  have  been  diversified,  through  sexual  selection, 
for  the  sake  of  ornament.  Here,  as  both  sexes  are  equally  well 
protected,  we  clearly  see  that  the  females  have  not  been  pre- 
vented by  natural  selection  from  inheriting  the  colors  of  their 
male  parents;  so  that  we  must  look  to  the  law  of  sexually-limited 
transmission. 

In  all  parts  of  the  world  both  sexes  of  many  soft-billed  birds, 
especially  those  which  frequent  reeds  or  sedges,  are  obscurely 
colored.  No  doubt  if  their  colors  had  been  brilliant,  they 
would  have  been  much  more  conspicuous  to  their  enemies;  but 
whether  their  dull  tints  have  been  specially  gained  for  the  sake 
of  protection  seems,  as  far  as  I  can  judge,  rather  doubtful.  It 
is  still  more  doubtful  whether  such  dull  tints  can  have  been 
gained  for  the  sake  of  ornament.  We  must,  however,  bear  in  mind 
that  male  birds,  though  dull-colored,  often  differ  much  from 
their  females  (as  with  the  common  sparrow),  and  this  leads  to 
the  belief  that  such  colors  have  been  gained  through  sexual  selec- 
tion, from  being  attractive.  Many  of  the  soft-billed  birds  are 
songsters;  and  a  discussion  in  a  former  chapter  should  not  be 
forgotten,  in  which  it  was  shown  that  the  best  songsters  are 
rarely  ornamented  with  bright  tints.  It  would  appear  that  female 
birds,  as  a  general  rule,  have  selected  their  mates  either  for  their 
sweet  voices  or  gay  colors,  but  not  for  both  charms  combined. 
Some  species,  which  are  manifestly  colored  for  the  sake  of  pro- 
tection, such  as  the  jack-snipe,  woodcock,  and  night-jar,  are  like- 
wise marked  and  shaded,  according  to  our  standard  of  taste,  with 
extreme  elegance.  In  such  cases  we  may  conclude  that  both 
natural  and  sexual  selection  have  acted  conjointly  for  protection 
and  ornament.  Whether  any  bird  exists  which  does  not  possess 
some  special  attraction,  by  which  to  charm  the  opposite  sex, 
may  be  doubted.  When  both  sexes  are  so  obscurely  colored  that 
it  would  be  rash  to  assume  the  agency  of  sexual  selection,  and 
when  no  direct  evidence  can  be  advanced  showing  that  such 
colors  serve  as  a  protection,  it  is  best  to  own  complete  ignorance 
of  the  cause,  or,  which  comes  to  nearly  the  same  thing,  to  at- 
tribute the  result  to  the  direct  action  of  the  conditions  of  life. 

Both  sexes  of  many  birds  are  conspicuously,  though  not  bril- 
liantly colored,  such  as  the  numerous  black,  white,  or  piebald 
species;  and  these  colors  are  probably  the  result  of  sexual  selec- 
tion.   With  the  common  blackbird,  capercailzie,  blackcock,  black 


BIRDS— CONSPICUOUS  COLORS.  487 

scoter-duck  (Oidemia),  and  even  with  one  of  the  birds  of  paradise 
(Lophorina  atra),  the  males  alone  are  black,  whilst  the  females 
are  brown  or  mottled;  and  there  can  hardly  be  a  doubt  that 
blackness  in  these  cases  has  been  a  sexually  selected  character. 
Therefore  it  is  in  some  degree  probable  that  the  complete  or 
partial  blackness  of  both  sexes  in  such  birds  as  crows,  certain 
cockatoos,  storks,  and  swans,  and  many  marine  birds,  is  likewise 
the  result  of  sexual  selection,  accompanied  by  equal  transmission 
to  both  sexes;  for  blackness  can  hardly  serve  in  any  case  as  a 
protection.  With  several  birds,  in  which  the  male  alone  is  black, 
and  in  others  in  which  both  sexes  are  black,  the  beak  or  skin 
about  the  head  is  brightly  colored,  and  the  contrast  thus  afforded 
adds  much  to  their  beauty;  we  see  this  in  the  bright  yellow 
beak  of  the  male  blackbird,  in  the  crimson  skin  over  the  eyes  of 
the  black-cock  and  capercailzie,  in  the  brightly  and  variously  col- 
ored beak  of  the  scoter-drake  (Oidemia),  in  the  red  beak  of  the 
chough  (Corvus  graculus,  Linn.),  of  the  black  swan,  and  the 
black  stork.  This  leads  me  to  remark  that  it  is  not  incredible 
that  toucans  may  owe  the  enormous  size  of  their  beaks  to  sexual 
selection  for  the  sake  of  displaying  the  diversified  and  vivid 
stripes  of  color,  with  which  these  organs  are  ornamented.^^  The 
naked  skin,  also,  at  the  base  of  the  beak  and  round  the  eyes  is 
likewise  often  brilliantly  colored;  and  Mr.  Gould,  in  speaking  of 
one  species,^^  says  that  the  colors  of  the  beak  "are  doubtless  in 
"the  finest  and  most  brilliant  state  during  the  time  of  pairing." 
There  is  no  greater  improbability  that  toucans  should  be  encum- 
bered with  immense  beaks,  though  rendered  as  light  as  possible 
by  their  cancellated  structure,  for  the  display  of  fine  colors  (an 
object  falsely  appearing  to  us  unimportant),  than  that  the  male 
Argus  pheasant  and  some  other  birds  should  be  encumbered  with 
plumes  so  long  as  to  impede  their  flight. 

In  the  same  manner,  as  the  males  alone  of  various  species  are 
black,  the  females   being  dull-colored;     so   in  a  few   cases  the 

SI  No  satisfactory  explanation  has  ever  been  offered  of  the  immense 
size,  and  still  less  of  the  bright  colors,  of  the  toucan's  beak.  Mr. 
Bates  ('The  Naturalist  on  the  Amazons,'  vol.  ii.  1863,  p.  341)  states  that 
they  use  their  beaks  for  reaching"  fruit  at  the  extreme  tips  of  the 
branches;  and  likewise,  as  stated  by  other  authors,  for  extracting  eggs 
and  young  birds  from  the  nests  of  other  birds.  But,  as  Mr.  Bates 
admits,  the  beak  "can  scarcely  be  considered  a  very  perfectly-formed 
"instrument  for  the  end  to  which  it  is  applied."  The  great  bulk  of  the 
beak,  as  shown  by  its  breadth,  depth,  as  well  as  length,  is  not  intelli- 
gible on  the  view,  that  It  serves  merely  as  an  organ  of  prehension. 
Mr.  Belt  believes  ('The  Naturalist  in  Nicaragua,'  p.  197),  that  the  prin- 
cipal use  of  the  beak  is  as  a  defense  against  enemies,  especially  to  the 
female  whilst  nesting  in  a  hole  in  a  tree. 

^*  Ramphastos  carinatus,  Gould's  'Monograph  of  Ramphastidae.' 


488  THE    DESCENT  OF  MAN. 

males  alone  are  either  wholly  or  partially  white,  as  with  the 
several  bell-birds  of  South  America  (Chasmorhynchus),  the  Ant- 
arctic goose  (Bernicla  antarctica),  the  silver-pheasant,  &c.,  whilst 
the  females  are  brown  or  obscurely  mottled.  Therefore,  on  the 
same  principle  as  before,  it  is  probable  that  both  sexes  of  many 
birds,  such  as  white  cockatoos,  several  egrets  with  their  beautiful 
plumes,  certain  ibises,  gulls,  terns,  &c.,  have  acquired  their  more 
or  less  completely  white  plumage  through  sexual  selection.  In 
some  of  these  cases  the  plumage  becomes  white  only  at  maturity. 
This  is  the  case  with  certain  gannets,  tropic-birds,  &c.,  and  with 
the  snow-goose  (Anser  hyperboreus).  As  the  latter  breeds  on  the 
"barren  grounds,"  when  not  covered  with  snow,  and  as  it  mi- 
grates southward  during  the  winter,  there  is  no  reason  to  suppose 
that  its  snow-white  adult  plumage  serves  as  a  protection.  In  the 
Anastomus  oscitans,  we  have  still  better  evidence  that  the  white 
plumage  is  a  nuptial  character,  for  it  is  developed  only  during 
the  summer;  the  young  in  their  immature  state,  and  the  adults 
in  their  winter  dress,  being  gray  and  black.  With  many  kinds 
of  gulls  (Larus),  the  head  and  neck  become  pure  white  during 
the  summer,  being  gray  or  mottled  during  the  winter  and  in  the 
young  state.  On  the  other  hand,  with  the  smaller  gulls,  or  sea- 
mews  (Gavia),  and  with  some  terns  (Sterna),  exactly  the  reverse 
occurs;  for  the  heads  of  the  young  birds  during  the  first  year, 
and  of  the  adults  during  the  winter,  are  either  pure  white,  or 
much  paler  colored  than  during  the  breeding-season.  These  latter 
cases  offer  another  instance  of  the  capricious  manner  in  which 
Bexual  selection  appears  often  to  have  acted.^^ 

That  aquatic  birds  have  acquired  a  white  plumage  so  much 
oftener  than  terrestrial  birds,  probably  depends  on  their  large 
size  and  strong  powers  of  flight,  so  that  they  can  easily  defend 
themselves  or  escape  from  birds  of  prey,  to  which  moreover  they 
are  not  much  exposed.  Consequently,  sexual  selection  has  not 
here  been  interfered  with  or  guided  for  the  sake  of  protection. 
No  doubt  with  birds  which  roam  over  the  open  ocean,  the  males 
and  females  could  find  each  other  much  more  easily,  when  made 
conspicuous  either  by  being  perfectly  white  or  intensely  black; 
so  that  these  colors  may  possibly  serve  the  same  end  as  the 
call-notes  o£  many  land-birds.^*    A  white  or  black  bird  when  it 


ss  On  Larus,  Gavia,  and  Sterna,  see  Macg-illivray,  'Hist.  Brit.  Birds,* 
vol,  v.  pp.  515,  584,  626.  On  the  Anser  hyperboreus,  Aubudon,  'Ornith. 
Biography,*  vol.  iv.  p.  562.  On  the  Anastomus,  Mr.  Blyth,  in  'Ibis,* 
1867,  'p.  173. 

5*  It  may  be  noticed  that  with  vultures,  which  roam  far  and  wide 
high  in  the  air,  like  marine  birds  over  the  ocean,  three  or  four  species 
are  almost  wholly  or  largely  white,  and  that  many  others  are  black. 
So  that  here  again  conspicuous  colors  may  possibly  aid  th/?  sexes  in 
finding  each  other  during  the  breeding  seassii,. 


BIRDS— CONSPICUOUS  COLORS.  489 

discovers  and  flies  down  to  a  carcass  floating  on  the  sea  or  cast 
up  on  the  beach,  will  be  seen  from  a  great  distance,  and  will 
guide  other  birds  of  the  same  and  other  species,  to  the  prey;  but 
as  this  would  be  a  disadvantage  to  the  first  finders,  the  individuals 
which  were  the  whitest  or  blackest  would  not  thus  procure  more 
food  than  the  less  strongly  colored  individuals.  Hence  conspicu- 
ous colors  cannot  have  been  gradually  acquired  for  this  purpose 
through  natural  selection. 

As  sexual  selection  depends  on  so  fluctuating  an  element  as 
taste,  we  can  understand  how  it  is  that,  within  the  same  group 
of  birds  having  nearly  the  same  habits,  there  should  exist  white 
or  nearly  white,  as  well  as  black,  or  nearly  black  species, — for 
instance,  both  white  and  black  cockatoos,  storks,  ibises,  swans, 
terns,  and  petrels.  Piebald  birds  likewise  sometimes  occur  in 
the  same  groups  together  with  black  and  white  species;  for 
instance,  the  black-necked  swan,  certain  terns,  and  the  common 
magpie.  That  a  strong  contrast  in  color  is  agreeable  to  birds, 
we  may  conclude  by  looking  through  any  large  collection,  for 
the  sexes  often  differ  from  each  other  in  the  male  having  the  pale 
parts  of  a  purer  white,  and  the  variously  colored  dark  parts  of 
still  darker  tints  than  the  female. 

It  would  even  appear  that  mere  novelty,  or  slight  changes  for 
the  sake  of  change,  have  sometimes  acted  on  female  birds  as  a 
charm,  like  changes  of  fashion  with  us.  Thus  the  males  of  some 
parrots  can  hardly  be  said  to  be  more  beautiful  than  the  females, 
at  least  according  to  our  taste,  but  they  differ  in  such  points, 
as  in  having  a  rose-colored  collar  instead  of  "a  bright  emeraldine 
"narrow  green  collar;"  or  in  the  male  having  a  black  collar  in- 
stead of  "a  yellow  demi-collar  in  front,"  with  a  pale  roseate  in- 
stead of  a  plum-blue  head.^^  As  so  many  male  birds  have  elon- 
gated tail-feathers  or  elongated  crests  for  their  chief  ornarpent, 
the  shortened  tail,  formerly  described  in  the  male  of  a  humming- 
bird, and  the  shortened  crest  of  the  male  goosander,  seem  like 
one  of  the  many  changes  of  fashion  which  we  admire  in  our  own 
dresses. 

Some  members  of  the  heron  family  offer  a  still  more  curious 
case  of  novelty  in  coloring,  having,  as  it  appears,  been  appre- 
ciated for  the  sake  of  novelty.  The  young  of  the  Ardea  asha  are 
white,  the  adults  being  dark  slate  colored;  and  not  only  the 
young  of  these  two  species,  as  well  as  of  some  other  members  of 
Buphus  coromandus  are  white,  this  color  changing  into  a  rich 
golden-buff  during  the  breeding-season.  It  is  incredible  that  the 
young  of  these  two  species,  as  well  as  of  some  other  members  of 

S5  See  Jer(5on  on  the  genus  Palaeornis,  'Birds  of  India,'  vol.  i.  p. 
253-260. 


490  THE    DESCENT  OF  MAN. 

the  same  family/^  should  for  any  special  purpose,  have  been  ren- 
dered pure  white  and  thus  made  conspicuous  to  their  enemies; 
or  that  the  adults  of  one  of  these  two  species  should  have  been 
specially  rendered  white  during  the  winter  in  a  country  which  is 
never  covered  with  snow.  On  the  other  hand  we  have  good  rea- 
son to  believe  that  whiteness  has  been  gained  by  many  birds  as  a 
sexual  ornament.  We  may  therefore  conclude  that  some  early 
progenitor  of  the  Ardea  asha  and  the  Buphus  acquired  a  white 
plumage  for  nuptial  purposes,  and  transmitted  this  color  to  their 
young;  so  that  the  young  and  the  old  became  white  like  certain 
existing  egrets;  and  that  the  whiteness  was  afterwards  retained 
by  the  young,  whilst  it  was  exchanged  by  the  adults  for  more 
strongly-pronounced  tints.  But  if  we  could  look  still  further  back 
to  the  still  earlier  progenitors  of  these  two  species,  we  should 
probably  see  the  adults  dark-colored.  I  infer  that  this  would  be 
the  case,  from  the  analogy  of  many  other  birds,  which  are  dark 
whilst  young,  and  when  adult  are  white;  and  more  especially 
from  the  case  of  the  Ardea  gularis,  the  colors  of  which  are  the 
reverse  of  those  of  A.  asha,  for  the  young  are  dark-colorfed  and 
the  adults  white,  the  young  having  retained  a  former  state  of 
plumage.  It  appears  therefore  that,  during  a  long  line  of  descent, 
the  adult  progenitors  of  the  Ardea  asha,  the  Buphus,  and  of  some 
allies,  have  undergone  the  following  changes  of  color:  first,  a 
dark  shade;  secondly,  pure  white;  and  thirdly,  owing  to  another 
change  of  fashion  (if  I  may  so  express  myself),  their  present  slaty, 
reddish,  or  golden-buff  tints.  These  successive  changes  are  in- 
telligible only  on  the  principle  of  novelty  having  been  admired  by 
birds  for  its  own  sake. 

Several  writers  have  objected  to  the  whole  theory  of  sexual 
selection,  by  assuming  that  with  animals  and  savages  the  taste 
of  the  female  for  certain  colors  or  other  ornaments  would  not 
remain  constant  for  many  generations;  that  first  one  color  and 
then  another  would  be  admired,  and  consequently  that  no  per- 
manent effect  could  be  produced.  We  may  admit  that  taste  is 
fluctuating,  but  it  is  not  quite  arbitrary.  It  depends  much  on 
habit,  as  we  see  in  mankind;  and  we  may  infer  that  this  would 
hold  good  with  birds  and  other  animals.  Even  in  our  own  dress, 
the  general  character  lasts  long,  and  the  changes  are  to  a  certain 
extent  graduated.  Abundant  evidence  will  be  given  in  two  places 
in  a  future  chapter,  that  savages  of  many  races  have  admired  for 
many  generations  the  same  cicatrices  on  the  skin,  the  same  bide- 


ts The  young  of  Ardea  rufescens  and  A.  coerulea  of  the  U.  States 
are  likewise  white,  the  adults  being  colored  in  accordance  with  their 
specific  names.  Audubon  ('Ornith.  Biography,'  vol.  iii.  p.  416;  vol.  iv. 
p.  58)  seems  rather  pleased  at  the  thought  that  this  remarkable  change 
of  plumage  will  greatly  "disconcert  the  systematists." 


BIRDS— SUMMARY.  491 

oiisly  perforated  lips,  nostrils,  or  ears,  distorted  heads,  &c.;  and 
these  deformities  present  some  analogy  to  the  natural  ornaments 
of  various  animals.  Nevertheless,  with  savageis  such  fashions 
do  not  endure  for  ever,  as  we  may  infer  from  the  differences  in 
this  respect  between  allied  tribes  on  the  same  continent.  So 
again  the  raisers  of  fancy  animals  certainly  have  admired  for 
many  generations  and  still  admire  the  same  breeds;  they  earnest- 
ly desire  slight  changes,  which  are  considered  as  improvements, 
but  any  great  or  sudden  change  is  looked  at  as  the  greatest  blem- 
ish. With  birds  in  a  state  of  nature  we  have  no  reason  to  suppose 
that  they  would  admire  an  entirely  new  style  of  coloration,  even 
if  great  and  sudden  variations  often  occurred,  which  is  far  from 
being  the  case.  We  know  that  dovecot  pigeons  do  not  willingly 
associate  with  the  variously  colored  fancy  breeds;  that  albino 
birds  do  not  commonly  get  partners  in  marriage;  and  that  the 
black  ravens  of  the  Feroe  Islands  chase  away  their  piebald  breth- 
ren. But  this  dislike  of  a  sudden  change  would  not  preclude  their 
appreciating  slight  changes,  any  more  than  it  does  in  the  case  of 
man.  Hence  with  respect  to  taste,  which  depends  on  many  ele- 
ments, but  partly  on  habit  and  partly  on  a  love  of  novelty,  there 
seems  no  improbability  in  animals  admiring  for  a  very  long  period 
the  same  general  style  of  ornamentation  or  other  attrac- 
tions, and  yet  appreciating  slight  changes  in  colors,  form,  or 
sound. 

Summary  of  the  Four  Chapters  on  Birds. — Most  male  birds  are 
highly  pugnacious  during  the  breeding-season,  and  some  possess 
weapons  adapted  for  fighting  with  their  rivals.  But  the  most 
pugnacious  and  the  best  armed  males  rarely  or  never  depend  for 
success  solely  on  their  power  to  drive  away  or  kill  their  rivals, 
but  have  special  means  for  charming  the  female.  With  some  it 
is  the  power  of  song,  or  of  giving  forth  strange  cries,  or  Instru- 
mental music,  and  the  males  in  consequence  differ  from  the  fe- 
males in  their  vocal  organs,  or  in  the  structure  of  certain  feathers. 
From  the  curiously  diversified  means  for  producing  various 
sounds,  we  gain  a  high  idea  of  the  importance  of  this  means  of 
courtship.  Many  birds  endeavor  to  charm  the  females  by  love- 
dances  or  antics,  performed  on  the  ground  or  in  the  air,  and  some- 
times at  prepared  places.  But  ornaments  of  many  kinds,  the 
most  brilliant  tints,  combs  and  wattles,  beautiful  plumes,  elon- 
gated feathers,  top-knots,  and  so  forth,  are  by  far  the  commonest 
means.  In  some  cases  mere  novelty  appears  to  have  acted  as  a 
charm.  The  ornaments  of  the  males  must  be  highly  important  to 
them,  for  they  have  been  acquired  in  not  a  few  cases  at  the  cost  of 
increased  danger  from  enemies,  and  even  at  some  loss  of  power 
in  fighting  with  their  rivals.  The  males  of  very  many  species 
do  not  assume  their  ornamental  dress  until  they  arrive  at  ma- 


492  THE  DESCENT  OF  MAN. 

turity,  or  they  assume  it  only  during  the  breeding-season,  or 
the  tints  then  become  more  vivid.  Certain  ornamental  appen- 
dages become  enlarged,  turgid,  and  brightly  colored  during  the 
act  of  courtship.  The  males  display  their  charms  with  elaborate 
care  and  to  the  best  effect;  and  this  is  done  in  the  presence  of 
the  females.  The  courtship  is  sometimes  a  prolonged  affair,  and 
many  males  and  females  congregate  at  an  appointed  place.  To 
suppose  that  the  females  do  not  appreciate  the  beauty  of  the 
males,  is  to  admit  that  their  splendid  decorations,  all  their  pomp 
and  display,  are  useless;  and  this  is  incredible.  Birds  have  fine 
powers  of  discrimination,  and  in  some  few  instances  it  can  be 
shown  that  they  have  a  taste  for  the  beautiful.  The  females, 
moreover,  are  known  occasionally  to  exhibit  a  marked  preference 
or  antipathy  for  certain  individual  males. 

If  it  be  admitted  that  the  females  prefer,  or  are  unconsciously 
excited  by  the  more  beautiful  males,  then  the  males  would  slowly 
but  surely  be  rendered  more  and  more  attractive  through  sexual 
selection.     That  it  is  this  sex  which  has  been  chiefly  modified, 
we  may  infer  from  the  fact  that,  in  almost  every  genus  where 
the  sexes  differ,  the  males  differ  much  more  from  one  another 
than  do  the  females;    this  is  well  shown  in  certain  closely- allied 
representative  species,  in  which  the  females  can  hardly  be  distin- 
guished, whilst  the  males  are  quite  distinct.  Birds  in  a  state  of  na- 
ture offer  individual  differences  which  would  amply  suffice  for  the 
work  of  sexual  selection;  but  we  have  seen  that  they  occasionally 
present  more  strongly-marked  variations  which  recur  so  frequent- 
ly that  they  would  immediately  be  fixed,  if  they  served  to  allure 
the  female.  The  laws  of  variation  must  determine  the  nature  of  the 
initial  changes,  and  will  have  largely  influenced  the  final  result. 
The  gradations,  which  may  be  observed  between  the  males  of 
allied  species,  indicate  the  nature  of  the  steps  through  which 
they  have  passed.    They  explain  also  in  the  most  interesting  man- 
ner how  certain  characters  have  originated,  such  as  the  indented 
ocelli  on  the  tail-feathers  of  the  peacock,  and  the  ball  and  socket 
ocelli  on  the  wing-feathers  of  the  Argus  pheasant.    It  is  evident 
that  the  brilliant  colors,  top-knots,  fine  plumes,  &c.,  of  many 
male  birds  cannot  have  been  acquired  as  a  protection;    indeed, 
they  sometimes  lead  to  danger.      That  they  are  not  due  to  the 
direct  and  definite  action  of  the  conditions  of  life,  we  may  feel 
assured,  because  the  females  have  been  exposed  to  the  same  con- 
ditions, and  yet  often  differ  from  the  males  to  an  extreme  degree. 
Although  it  is  probable  that  changed  conditions  acting  during  a 
lengthened  period  have  in  some  cases  produced  a  definite  effect 
on  both  sexes,  or  sometimes  on  one  sex  alone,  the  more  important 
result  will  have  been  an  increased  tendency  to  vary  or  to  present 
more  strongly  marked  individual  differences;    and  such  differ- 


BIRDS— SUMMARY.  493 

ences  will  have  afforded  an  excellent  ground-work  for  the  action 
of  sexual  selection. 

The  laws  of  inheritance,  irrespectively  of  selection,  appear  to 
have  determined  whether  the  characters  acquired  by  the  males 
for  the  sake  of  ornament,  for  producing  various  sounds,  and  for 
fighting  together,  have  been  transmitted  to  the  males  alone  or  to 
both  sexes,  either  permanently,  or  periodically  during  certain 
seasons  of  the  year.  Why  various  characters  should  have  been 
transmitted  sometimes  in  one  way  and  sometimes  in  another,  is 
not  in  most  cases  known;  but  the  period  of  variability  seems 
often  to  have  been  the  determining  cause.  When  the  two  sexes 
have  inherited  all  characters  in  common  they  necessarily  resemble 
each  other;  but  as  the  successive  variations  may  be  differently 
transmitted,  every  possible  gradation  may  be  found,  even  within 
the  same  genus,  from  the  closest  similarity  to  the  widest  dis- 
similarity between  the  sexes.  With  many  closely-allied  species, 
following  nearly  the  same  habits  of  life,  the  males  have  come  to 
differ  from  each  other  chiefly  through  the  action  of  sexual  selec- 
tion; whilst  the  females  have  come  to  differ  chiefly  from  partak- 
ing more  or  less  of  the  characters  thus  acquired  by  the  males. 
The  effects,  moreover,  of  the  definite  action  of  the  conditions  of 
life,  will  not  have  been  masked  in  the  females,  as  in  the  males, 
by  the  accumulation  through  sexual  selection  of  strongly-pro- 
nounced colors  and  other  ornaments.  The  individuals  of  both 
sexes,  however  affected,  will  have  been  kept  at  each  successive 
period  nearly  uniform  by  the  free  intercrossing  of  many  individ- 
uals. 

With  species,  in  which  the  sexes  differ  in  color,  it  is  possible 
or  probable  that  some  of  the  successive  variations  often  tended 
to  be  transmitted  equally  to  both  sexes;  but  that  when  this  oc- 
curred the  females  were  prevented  from  acquiring  the  bright 
colors  of  the  males,  by  the  destruction  which  they  suffered  dur- 
ing incubation.  There  is  no  evidence  that  it  is  possible  "by  natural 
selection  to  convert  one  form  of  transmission  into  another.  But 
there  would  not  be  the  least  difficulty  in  rendering  a  female  dull- 
colored,  the  male  being  still  kept  bright-colored,  by  the  selection 
of  successive  variations,  which  were  from  the  first  limited  in 
their  transmission  to  the  same  sex.  Whether  the  females  of  many 
species  have  actually  been  thus  modified,  must  at  present  remain 
doubtful.  When,  through  the  law  of  the  equal  transmission  of 
characters  to  both  sexes,  the  females  were  rendered  as  conspicu- 
ously colored  as  the  males,  their  instincts  appear  often  to  have 
been  modified  so  that  they  were  led  to  build  domed  or  concealed 
nests. 

In  one  small  and  curious  class  of  cases  the  characters  and 
habits  of  the  two  sexes  have  been  completely  transposed,  for  the 
females  are  larger,  stronger,  more  vociferous  and  brighter  col- 


494  THE  DESCENT  OF  MAN. 

ored  than  the  males.  They  have,  also,  become  so  quarrelsome 
that  they  often  fight  together  for  the  possession  of  the  males,  like 
the  males  of  other  pugnacious  species  for  the  possession  of  the 
females.  If,  as  seems  probable,  such  females  habitually  drive 
away  their  rivals,  and  by  the  display  of  their  bright  colors  or 
other  charms  endeavor  to  attract  the  males,  we  can  understand 
how  it  is  that  they  have  gradually  been  rendered,  by  sexual  selec- 
tion and  sexually-limited  transmission,  more  beautiful  than  the 
males — the  latter  being  left  unmodified  or  only  slightly  modi- 
fied. 

Whenever  the  law  of  inheritance  at  corresponding  ages  prevails, 
but  not  that  of  sexually-limited  transmission,  then  if  the  parents 
vary  late  in  life — and  we  know  that  this  constantly  occurs  with 
our  poultry,  and  occasionally  with  other  birds — the  young  will  be 
left  unaffected,  whilst  the  adults  of  both  sexes  will  be  modified. 
If  both  these  laws  of  inheritance  prevail  and  either  sex  varies 
late  in  life,  that  sex  alone  will  be  modified,  the  other  sex  and  the 
young  being  unaffected.  When  variations  in  brightness  or  in 
other  conspicuous  characters  occur  early  in  life,  as  no  doubt  often 
happens,  they  will  not  be  acted  on  through  sexual  selection  until 
the  period  of  reproduction  arrives;  consequently  if  dangerous  to 
the  young,  they  will  be  eliminated  through  natural  selection. 
Thus  we  can  understand  how  it  is  that  variations  arising  late  in 
life  have  so  often  been  preserved  for  the  ornamentation  of  the 
males;  the  females  and  the  young  being  left  almost  unaffected,  and 
therefore  like  each  other.  With  species  having  a  distinct  sum- 
mer and  winter  plumage,  the  males  of  which  either  resemble  or 
differ  from  the  females  during  both  seasons  or  during  the  summer 
alone,  the  degrees  and  kinds  of  resemblance  between  the  young 
and  the  old  are  exceedingly  complex;  and  this  complexity  ap- 
parently depends  on  characters,  first  acquired  by  the  males,  being 
transmitted  in  various  ways  and  degrees,  as  limited  by  age,  sex, 
and  season. 

As  the  young  of  so  many  species  have  been  but  little  modified 
in  color  and  in  other  ornaments,  we  are  enabled  to  form  some 
judgment  with  respect  to  the  plumage  of  their  early  progenitors; 
and  we  may  infer  that  the  beauty  of  our  existing  species,  if  we 
look  to  the  whole  class,  has  been  largely  increased  since  that 
period,  of  which  the  immature  plumage  gives  us  an  indirect  rec- 
ord. Many  birds,  especially  those  which  live  much  on  the  ground, 
have  undoubtedly  been  obscurely  colored  for  the  sake  of  protec- 
tion. In  some  instances  the  upper  exposed  surface  of  the  plumage 
has  been  thus  colored  in  both  sexes,  whilst  the  lower  surface  in 
the  males  alone  has  been  variously  ornamented  through  sexual 
selection.  Finally,  from  the  facts  given  in  these  four  chapters, 
we  may  conclude  that  weapons  for  battle,  organs  for  producing 
sound,  ornaments  of  many  kinds,  bright  and  conspicuous  colors. 


BIRDS— SUMMARY.  495 

have  generally  been  acquired  hy  the  males  through  variation  ana 
sexual  selection,  and  have  been  transmitted  m  various  ways  ac- 
cording to  the  several  laws  of  inheritance — the  females  and  the 
young  being  left  comparatively  but  little  modified.^^ 

"  I  am  greatly  indebted  to  the  kindness  of  Mr.  Sclater  for  having 
looked  over  these  four  chapters  on  birds,  and  the  two  following  ones 
on  mammals.  In  this  way  I  have  been  saved  from  making  mistakes 
about  the  names  of  the  species,  and  from  stating  anything  as  a  fact 
which  is  known  to  this  distinguished  naturalist  to  be  erroneous.  But 
of  course  he  is  not  at  all  answerable  for  the  accuracy  of  the  state- 
ments quoted  by  me  from  various  authorities. 


496  THE  DESCENT  OF  MAN. 


CHAPTER  XVII. 

SECONDARY  SEXUAL.  CHARACTERS  OF  MAMMALS. 

The  law  of  battle— Special  weapons,  confined  to  the  males— Cause  of 
absence  of  weapons  In  the  female— Weapons  common  to  both  sexes, 
yet  primarily  acquired  by  the  male — Other  uses  of  such  weapons— 
Their  high  importance — Greater  size  of  the  male — Means  of  defense 
— On  the  preference  shown  by  either  sex  in  the  pairing  of  quadru- 
peds. 

With  mammals  the  male  appears  to  win  the  female  much  more 
through  the  law  of  battle  than  through  the  display  of  his 
charms.  The  most  timid  animals,  not  provided  with  any  special 
weapons  for  fighting,  engage  in  desperate  conflicts  during  the 
season  of  love.  Two  male  hares  have  been  seen  to  fight  together 
until  one  was  killed;  male  moles  often  fight,  and  sometimes 
with  fatal  results;  male  squirrels  engage  in  frequent  contests, 
"and  often  wound  each  other  severely;"  as  do  male  beavers,  so 
that  "hardly  a  skin  is  without  scars."^  I  observed  the  same  fact 
with  the  hides  of  the  guanacoes  in  Patagonia;  and  on  one  oc- 
casion several  were  so  absorbed  in  fighting  that  they  fearlessly 
rushed  close  by  me.  Livingstone  speaks  of  the  males  of  the 
many  animals  in  Southern  Africa  as  almost  invariably  showing 
the  scars  received  in  former  contests. 

The  law  of  battle  prevails  with  aquatic  as  with  terrestrial  mam- 
mals. It  is  notorious  how  desperately  male  seals  fight,  both  with 
their  teeth  and  claws,  during  the  breeding-season;  and  their 
hides  are  likewise  often  covered  with  scars.  Male  sperm-whales 
are  very  jealous  at  this  season;  and  in  their  battles  ''they  often 
"lock  their  jaws  together,  and  turn  on  their  sides  and  twist 
"about;"  so  that  their  lower  jaws  often  become  distorted.^' 


1  See  Waterton's  account  of  two  hares  fighting,  'Zoologist,'  vol.  i. 
1843,  p.  211.  On  moles,  Bell,  'Hist,  of  British  Quadrupeds,'  1st  edit.  p. 
100. '  On  squirrels,  Audubon  and  Bachman,  'Viviparous  Quadrupeds  of 
N.  America,'  1846.  p.  269.  On  beavers,  Mr.  A.  H.  Green,  in  'Journal  of 
Lin.  Soc.  Zoolog.'  vol.  x.  1869,  p.  362. 

-  On  the  battles  of  seals,  see  Capt.  C.  Abbott  in  'Proc.  Zool.  Soc' 
1868,  p.  191;  also  Mr.  R.  Brown,  ibid.  1868,  p.  436;  also  L.  Lloyd,  'Game 


MAMMALS— LAW  OF  BATTLE.  49? 

All  male  animals  which  are  furnished  with  special  weapons 
for  fighting,  are  well  known  to  engage  in  fierce  battles.  The  cour- 
age and  the  desperate  conflicts  of  stags  have  often  been  described; 
their  skeletons  have  been  found  in  various  parts  of  the  world,  with 
the  horns  inextricably  locked  together,  showing  how  miserably 
the  victor  and  vanquished  had  perished."  No  animal  in  the  world 
is  so  dangerous  as  an  elephant  in  must.  Lord  Tankerville  has 
given  me  a  graphic  description  of  the  battles  between  the  wild 
bulls  in  Chillingham  Park,  the  descendants,  degenerated  in  size 
but  not  in  courage,  of  the  gigantic  Bos  primigenius.  In  1861  sev- 
eral contended  for  mastery;  and  it  was  observed  that  two  of  the 
younger  bulls  attacked  in  concert  the  old  leader  of  the  herd, 
overthrew  and  disabled  him,  so  that  he  was  believed  by  the 
keepers  to  be  lying  mortally  wounded  in  a  neighboring  wood. 
But  a  few  days  afterwards  one  of  the  young  bulls  approached  the 
Wood  alone;  and  then  the  "monarch  of  the  chase,"  who  had  been 
lashing  himself  up  for  vengeance,  came  out  and,  in  a  short  time, 
killed  his  antagonist.  He  then  quietly  joined  the  herd,  and  long 
held  undisputed  sway.  Admiral  Sir  J.  B.  Sulivan  informs  me 
that,  when  he  lived  in  the  Falkland  Islands,  he  imported  a  young 
English  stallion,  which  frequented  the  hills  near  Port  William 
with  eight  mares.  On  these  hills  there  were  two  wild  stallions, 
each  with  a  small  tr'oop  of  mares;  "and  it  is  certain  that  these 
"stallions  would  never  have  approached  each  other  without  fight- 
"ing.  Both  had  tried  singly  to  fight  the  English  horse  and  drive 
"away  his  mares,  but  had  failed.  One  day  they  came  in  together 
"and  attacked  him.  This  was  seen  by  the  capitan  who  had  charge 
"of  the  horses,  and  who,  on  riding  to  the  spot,  found  one  of  the 
"two  stallions  engaged  with  the  English  horse,  whilst  the  other 
"was  driving  away  the  mares,  and  had  already  separated  four  from 
"the  rest.  The  capitan  settled  the  matter  by  driving  the  whole 
"party  into  the  corral,  for  the  wild  stallions  would  not  leave  the 
"mares." 

Male  animals  which  are  provided  with  efficient  cutting  or  tear- 
ing teeth  for  the  ordinary  purposes  of  life,  such  as  the  carnivora, 
insectivora,  and  rodents,  are  seldom  furnished  with  weapons 
especially  adapted  for  fighting  with  their  rivals.  The  case  is 
very  different  with  the  males  of  many  other  animals.  We  see 
this  in  the  horns  of  stags  and  of  certain  kinds  of  antelopes  in 

Birds  of  Sweden,'  1867,  p.  412;  also  Pennant.    On  the  sperm-whale,  see 
Mr.  J.  H.  Thompson,  in  'Proc.  Zool.  Soc'  1867,  p.  246. 

3  See  Scrope  (Art  of  Deer-stalking-,'  p.  17)  on  the  locking-  of  the 
horns  with  the  Cervus  elaphus.  Richardson,  in  'Fauna  Bor.  Ameri- 
cana,' 1829,  p.  252,  says  that  the  wapiti,  moose,  and  rein-deer  have 
been  found  thus  locked  together.  Sir  A.  Smith  found  at  the  Cape  of 
Good  Hope  the  skeletons  of  two  gnus  in  the  same  condition. 
S3 


49S  THE  DESCENT  OF  MAN. 

which  the  females  are  hornless.  With  many  animals  the  canine 
teeth  in  the  upper  or  lower  jaw,  or  in  both,  are  much  larger  in  the 
males  than  in  the  females,  or  are  absent  in  the  latter,  with  the 
exception  sometimes  of  a  hidden  rudiment.  Certain  antelopes, 
the  musk-deer,  camel,  horse,  boar,  various  apes,  seals,  and  the 
walrus,  offer  instances.  In  the  females  of  the  walrus  the  tusks 
are  sometimes  quite  absent.*  In  the  male  elephant  of  India  and 
in  the  male  dugong^  the  upper  incisors  form  offensive  weapons. 
In  the  male  narwhal  the  left  canine  alone  is  developed  into  the 
well-known,  spirally-twisted,  so-called  horn,  which  is  sometimes 
from  nine  to  ten  feet  in  length.  It  is  believed  that  the  males  use 
these  horns  for  fighting  together;  for  "an  unbroken  one  can  rarely 
"be  got  and  occasionally  one  may  be  found  with  the  point  of 
"another  jammed  into  the  broken  place."^  The  tooth  on  the  op- 
posite side  of  the  head  in  the  male  consists  of  a  rudiment  about 
ten  inches  in  length,  v^^hich  is  embedded  in  the  jaw;  but  some- 
times, though  rarely,  both  are  equally  developed  on  the  two  sides. 
In  the  female  both  are  always  rudimentary.  The  male  cachalot 
has  a  larger  head  than  that  of  the  female,  and  it  no  doubt  aids  him 
in  his  aquatic  battles.  Lastly,  the  adult  male  ornithorhynchus  is 
provided  with  a  remarkable  apparatus,  namely  a  spur  on  the  fore- 
leg, closely  resembling  the  poison-fang  of  a  venomous  snake;  but 
according  to  Harting,  the  secretion  from  the  gland  is  not  poison- 
ous and  on  the  leg  of  the  female  there  is  a  hollow,  apparently  for 
the  reception  of  the  spur.^ 

When  the  males  are  provided  with  weapons  which  in  the  fe- 
males are  absent,  there  can  hardly  be  a  doubt  that  these  serve 
for  fighting  with  other  males;  and  that  they  were  acquired 
through  sexual  selection,  and  were  transmitted  to  the  male  sex 
alone.  It  is  not  probable,  at  least  in  most  cases,  that  the  females 
have  been  prevented  from  acquiring  such  weapons,  on  account  of 
their  being  useless,  superfluous,  or  in  some  way  injurious.  On 
the  contrary,  as  they  are  often  used  by  the  males  for  various 
purposes,  more  especially  as  a  defense  against  their  enemies,  it 

4  Mr.  Lamont  ('Seasons  with  the  Sea-Horses,'  1861,  p.  143)  says  that 
a  good  tusk  of  the  male  walrus  vv^eig-hs  4  pounds,  and  is  longer  than 
that  of  the  female,  which  weighs  about  3  pounds.  The  males  are 
described  as  fighting  ferociously.  On  the  occasional  absence  of  the 
tusks  in  the  female,  see  Mr.  R.  Brown,  'Proc.  Zool.  Soc'  1868,  p.  429. 

5  Owen,  'Anatomy  of  Vetebrates,'  vol.  iii.  p.  283. 

6  Mr.  R.  Brown,  in  'Proc.  Zool.  Soc'  1S69,  p.  553.  See  Prof.  Turner 
in  Journal  of  'Anat.  and  Phys.'  1872,  p.  76,  on  the  homological  nature  o 
these  tusks.  Also  Mr.  J.  W.  Clarke  on  two  tusks  being  developed  i:. 
the  males   in  'Proc.  Zoolog.  Soc'  1871,  p.  42. 

7  Owen  on  the  cachalot  and  Ornithorhynchus,  ibid.  vol.  iii.  pp.  638, 
641.  Harting  is  quoted  by  Dr.  Zouteveen  in  the  Dutch  translat.  of 
this  work,  vol.  ii.  p.  292. 


MAMMALS— LAW  OF  BATTLE.  499 

is  a  surprising  fact  that  they  are  so  poorly  developed,  or  quite 
absent  in  the  females  of  so  many  animals.  With  female  deer 
the  development  during  each  recurrent  season  of  great  branching 
horns,  and  with  female  elephants  the  development  of  immense 
tusks,  would  be  a  great  waste  of  vital  power,  supposing  that  they 
were  of  no  use  to  the  females.  Consequently,  they  would  have 
tended  to  be  eliminated  in  the  female  through  natural  selection; 
that  is,  if  the  successive  variations  were  limited  in  their  transmis- 
sion to  the  female  sex,  for  otherwise  the  weapons  of  the  males 
would  have  been  injuriously  affected,  and  this  would  have  been  a 
greater  evil.  On  the  whole,  and  from  the  consideration  of  the  fol- 
lowing facts,  it  seems  probable  that  when  the  various  weapons  dif- 
fer in  the  two  sexes,  this  has  generally  depended  on  the  kind  of 
transmission  which  has  prevailed. 

As  the  reindeer  is  the  one  species  in  the  whole  family  of  Deer, 
in  which  the  female  is  furnished  with  horns,  though  they  are  some- 
what smaller,  thinner,  and  less  branched  than  in  the  male,  it  might 
naturally  be  thought  that,  at  least  in  this  case,  they  must-  be  of 
some  special  service  to  her.  The  female  retains  her  horns  from 
the  time  when  they  are  fully  developed,  namely,  in  September, 
throughout  the  winter  until  April  or  May,  when  she  brings  forth 
her  young.  Mr.  Crotch  made  particular  enquiries  for  me  in  Nor- 
way, and  it  appears  that  the  females  at  this  season  conceal  them- 
selves for  about  a  fortnight  in  order  to  bring  forth  their  young, 
and  then  reappear,  generally  hornless.  In  Novia  Scotia  however,  as 
I  hear  from  Mr.  H.  Reeks,  the  female  sometimes  retains  her  horns 
longer.  The  male  on  the  other  hand  casts  his  horns  much  earlier, 
towards  the  end  of  November.  As  both  sexes  have  the  same  re- 
quirements and  follow  the  same  habits  of  life,  and  as  the  male  is 
destitute  of  horns  during  the  winter,  it  is  improbable  that  they 
can  be  of  any  special  service  to  the  female  during  this  season, 
which  includes  the  larger  part  of  the  time  during  which  she  is 
horned.  Nor  is  it  probable  that  she  can  have  inherited  horns 
from  some  ancient  progenitor  of  the  family  of  deer,  for,  from  the 
fact  of  the  females  of  so  many  species  in  all  quarters  of  the  globe 
not  having  horns,  we  may  conclude  that  this  was  the  primordial 
character  of  the  group.^ 

The  horns  of  the  reindeer  are  developed  at  a  most  unusually 
early  age;  but  what  the  cause  of  this  may  be  is  not  known.  The 
effect  has  apparently  been  the  transference  of  the  horns  to  both 
sexes.  We  should  bear  in  mind  that  horns  are  always  transmitted 
through  the  female,  and  that  she  has  a  latent  capacity  for  their  de- 

8  On  the  structure  and  shedding-  of  the  horns  of  the  reindeer,  Hoff- 
berg,  'Amoenitates  Acad.'  vol.  iv.  1788,  p.  149.  See  Richardson,  'Fauna 
Bor.  Americana,'  p.  241,  in  regard  to  the  American  variety  or  species; 
also  Major  W.  Ross  King,  'The  Sportsman  in  Canada,'  1866,  p.  80. 


500  THE  DESCENT  OP  MAN. 

velopment,  as  we  see  in  old  or  diseased  females."  Moreover  the  fe- 
males of  some  other  species  of  deer  exhibit,  either  normally  or 
occasionally,  rudiments  of  horns;  thus  the  female  of  Cervulus  mos- 
chatus  has  "bristly  tufts,  ending  in  a  knob,  Instead  of  a  horn;" 
and  "in  most  specimens  of  the  female  wapiti  (Cervus  canadensis) 
"there  is  a  sharp  bony  protuberance  in  the  place  of  the  horn."^" 
From  these  several  considerations  we  may  conclude  that  the  pos- 
session of  fairly  well-developed  horns  by  the  female  reindeer,  is 
due  to  the  males  having  first  acquired  them  as  weapons  for  fight- 
ing with  other  males;  and  secondarily  to  their  development  from 
some  unknown  cause  at  an  unusually  early  age  in  the  males,  and 
their  consequent  transference  to  both  sexes. 

Turning  to  the  sheath-horned  ruminants:  with  antelopes  a 
graduated  series  can  be  formed,  beginning  with  species,  the  fe- 
males of  which  are  completely  destitute  of  horns — passing  on  to 
those  which  have  horns  so  small  as  to  be  amost  rudimentary,  (as 
with  the  Antilocapra  americana,  in  which  species  they  are  present 
in  only  one  out  of  four  or  five  females") — to  those  which  have 
fairly  developed  horns,  but  manifestly  smaller  and  thinner  than 
in  the  male  and  sometimes  of  a  different  shape,^- — and  ending 
with  those  in  which  both  sexes  have  horns  of  equal  size.  As  with 
the  reindeer,  so  with  antelopes  there  exists,  as  previously  shown, 
a  relation  between  the  period  of  the  development  of  the  horns 
and  their  transmission  to  one  or  both  sexes;  it  is  there- 
fore probable  that  their  presence  or  absence  in  the 
females  of  some  species,  and  their  more  or  less  perfect 
condition  in  the  females  of  other  species,  depends,  not 
on  their  being  of  any  special  use,  but  simply  in  inheritance. 
It  accords  with  this  view  that  even  in  the  same  restricted  genus 
both  sexes  of  some  species,  and  the  males  alone  of  others,  are 
thus  provided.  It  is  also  a  remarkable  fact  that,  although  the 
females  of  Antilope  bezoartica  are  normally  destitute  of  horns, 


®  Isidore  Geoffrey  St.  Hilaire,  'Essais  de  Zoolog.  Generale,'  1841,  p. 
513.  Other  masculine  characters,  besides  the  horns,  are  sometimea 
similarly  transferred  to  the  female;  thus  Mr.  Boner,  in  speaking  of  an 
old  female  chamois  ('Chamois  Hunting  in  the  Mountains  of  Ba- 
varia,' 1850,  2nd  edit.  p.  363),  says,  "not  only  was  the  head  veiry  male- 
"looking,  but  along  the  back  there  was  a  ridge  of  long  hair,  usually 
"to  be  found  only  in  bucks." 

10  On  the  Cervulus,  Dr.  Gray,  'Catalogue  of  Mammalia  in  the  Brit- 
ish Museum,'  part.  iii.  p.  220.  On  the  Cervus  canadensis  or  wapiti,  see 
Hon.  J.  D.  Caton,  'Ottawa  Acad,  of  Nat.  Sciences,'  May,  1868,  p.  9. 

"  I  am  indebted  to  Dr.  Canfield,  for  this  information,  see  also  his 
paper  in  'Proc,  Zoolog.  Soc'  1866,  p.  105. 

12  For  instance  the  horns  of  the  female  Ant.  euchore  resemble  those 
of  a  distinct  species,  viz.  the  Ant.  dorcas  var.  Corine,  see  Desraarest, 
•Mammalogie,'  p.  455. 


Mammals— LAW  of  battle.  501 

Mr.  Blyth  has  seen  no  less  than  three  females  thus  furnished;  and 
there  was  no  reason  to  suppose  that  they  were  old  or  diseased. 

In  all  the  wild  species  of  goats  and  sheep  the  horns  are  larger 
in  the  male  than  in  the  female,  and  are  sometimes  quite  ahsent  in 
the  latter."  In  several  domestic  breeds  of  these  two  animals,  the 
males  alone  are  furnished  with  horns;  and  in  some  breeds,  for  in- 
stance, in  the  sheep  of  North  Wales,  though  both  sexes  are 
properly  horned,  the  ewes  are  very  liable  to  be  hornless.  I  have 
been  informed  by  a  trustworthy  witness,  who  purposely  inspected 
a  flock  of  these  same  sheep  during  the  lambing  season,  that  the 
horns  at  birth  are  generally  more  fully  developed  in  the  male 
than  the  female.  Mr.  J.  Peel  crossed  his  Lonk  sheep,  both  sexes 
of  which  always  bear  horns,  with  hornless  Leicesters  and  hornless 
Shropshire  Downs;  and  the  result  was  that  the  male  offspring  had 
their  horns  considerably  reduced,  whilst  the  females  were  wholly 
destitute  of  them.  These  several  facts  indicate  that,  with  sheep, 
the  horns  are  a  much  less  firmly  fixed  character  in  the  females 
than  in  the  males;  and  this  leads  us  to  look  at  the  horns  as 
properly  of  masculine  origin. 

With  the  adult  musk-ox  (Ovibos  moschatus)  the  horns  of  the 
male  are  larger  than  those  of  the  female,  and  in  the  latter  the 
bases  do  not  touch."  In  regard  to  ordinary  cattle  Mr.  Blyth  re- 
marks: *'In  most  of  the  wild  bovine  animals  the  horns  are  both 
"longer  and  thicker  in  the  bull  than  in  the  cow,  and  in  the  cow- 
"banteng  (Bos  sondaicus)  the  horns  are  remarkably  small,  and 
"inclined  much  backwards.  In  the  domestic  races  of  cattle,  both 
"of  the  humped  and  humpless  types,  the  horns  are  short  and  thick 
"in  the  bull,  longer  and  more  slender  in  the  cow  and  ox;  and  in 
"the  Indian  buffalo,  they  are  shorter  and  thicker  in  the  bull, 
"longer  and  more  slender  in  the  cow.  In  the  wild  gaour  (B. 
"gaurus)  the  horns  are  mostly  both  longer  and  thicker  in  the 
"bull  than  in  the  cow."^^  Dr.  Forsyth  Major  also  informs  me  that 
a  fossil  skull,  believed  to  be  that  of  the  female  Bos  etruscus,  has 
been  found  in  the  Val  d'Arno,  which  is  wholly  without  horns.  In 
the  Rhinoceros  simus,  as  I  may  add,  the  horns  of  the  female  are 
generally  longer  but  less  powerful  than  in  the  male;  and  in  some 
other  species  of  rhinoceros  they  are  said  to  be  shorter  in  the  fe- 
male.'® From  these  various  facts  we  may  infer  as  probable  that 
horns  of  all  kinds,  even  when  they  are  equally  developed  in  the 
two  sexes,  were  primarily  acquired  by  the  male  in  order  to  con- 

^  Gray,  'Catalogue  Mamm.  Brit.  Mus.'  part  iii.  1852,  p.  160. 
1*  Richardson,  'Fauna  Bor.  Americana,'  p.  278. 
IS  'Land  and  Water,'  1867,  p.  346. 

-«  Sir  Andrew  Smith,  'Zoology  of  S.  Africa,'  pi.  xix.  Owen,  'Anatomy 
of  Vertebrates,'  vol.  iii.  p.  624. 

33 


502  'J-'HE  DESCENT  OF  MAN. 

quer  other  males,  and  have  been  transferred  more  or  less  com- 
pletely to  the  female. 

The  effects  of  castration  deserve  notice,  as  throwing  light  on  this 
same  point.  Stags  after  the  operation  never  renew  their  horns. 
The  male  reindeer,  however,  must  be  excepted,  as  after  castration 
he  does  renew  them.  This  fact  as  well  as  the  possession  of  horns  by 
both  sexes,  seem  at  first  to  prove  that  the  horns  in  this  species  do 
not  constitute  a  sexual  character;"  but  as  they  are  developed  at 
a  very  early  age,  before  the  sexes  differ  in  constitution,  it  is  not 
surprising  that  they  should  be  unaffected  by  castration,  even  if 
they  were  aboriginally  acquired  by  the  male.  With  sheep  both 
sexes  properly  bear  horns;  and  I  am  informed  that  with  Welsh 
sheep  the  horns  of  the  males  are  considerably  reduced  by  castra- 
tion; but  the  degree  depends  much  on  the  age  at  which  the  opera- 
tion is  performed,  as  is  likewise  the  case  with  other  animals. 
Merino  rams  have  large  horns,  whilst  the  ewes  "generally  speak- 
"ing  are  without  horns;"  and  in  this  breed  castration  seems  to  pro- 
duce a  somewhat  greater  effect,  so  that  if  performed  at  an  early 
age  the  horns  "remain  almost  undeveloped."^^  On  the  Guinea 
coast  there  is  a  breed  in  which  the  females  never  bear  horns,  and, 
as  Mr.  Winwood  Reade  informs  me,  the  rams  after  castration  are 
quite  destitute  of  them.  With  cattle,  the  horns  of  the  males  are 
much  altered  by  castration;  for  instead  of  being  short  and  thick, 
they  become  longer  than  those  of  the  cow,  but  otherwise  resemble 
them.  The  Antilope  bezoartica  offers  a  somewhat  analogous  case: 
the  males  have  long  straight  spiral  horns,  nearly  parallel  to  each 
other,  and  directed  backwards;  the  females  occasionally  bear 
horns,  but  these  when  present  are  of  a  very  different  shape,  for 
they  are  not  spiral,  and  spreading  widely,  bend  round  with  the 
points  forwards.  Now  it  is  a  remarkable  fact  that,  in  the  cas- 
trated male,  as  Mr.  Blyth  informs  me,  the  horns  are  of  the  same 
peculiar  shape  as  in  the  female,  but  longer  and  thicker.  If  we 
may  judge  from  analogy,  the  female  probably  shows  us,  in  these 
two  cases  of  cattle  and  the  antelope,  the  former  condition  of  the 
horns  in  some  early  progenitor  of  each  species.  But  why  castra- 
tion should  lead  to  the  reappearance  of  an  early  condition  of  the 
horns  cannot  be  explained  with  any  certainty.  Nevertheless,  it 
seems  probable,  that  in  nearly  the  same  manner  as  the  consti- 
tutional disturbance  in  the  offspring,  caused  by  a  cross  between 


"  This  is  the  conclusion  of  Seldlitz,  'Die  Darwinsche  Theorie,'  1871, 
p.  47. 

18  I  am  much  obliged  to  Prof.  Victor  Carus,  for  having  made  in- 
quiries for  me  in  Saxony  on  this  subject.  H.  von  Nathusius  ('Vieh- 
zucht,'  1872,  p.  64)  says  that  the  horns  of  sheep  castrated  at  an  early 
period,  either  altogether  disappear  or  remain  as  mere  rudiments;  but 
I  do  not  know  whether  he  refers  to  merinos  or  to  ordinary  breeds. 


MAMMALS— LAW  OF  BATTLE.  502 

two  distinct  species  or  races,  often  leads  to  the  reappearance  of 
long-lost  characters  ;^^  so  here,  the  disturbance  in  the  constitution 
of  the  individual,  resulting  from  castration,  produces  the  same 
effect. 

The  tusks  of  the  elephant,  in  the  different  species  or  races,  differ 
according  to  sex,  nearly  as  do  the  horns  of  ruminants.  In  India 
and  Malacca  the  males  alone  are  provided  with  well-developed 
tusks.  The  elephant  of  Ceylon  is  considered  hy  most  naturalists 
as  a  distinct  race,  but  by  some  as  a  distinct  species,  and  here  "not 
"one  in  a  hundred  is  found  with  tusks,  the  few  that  possess  them 
"being  exclusively  males."'-"  The  African  elephant  is  undoubtedly 
distinct,  and  the  female  has  large  well-developed  tusks,  though 
not  so  large  as  those  of  the  male. 

These  differences  in  the  tusks  of  the  several  races  and  species 
of  elephants — the  gr^at  variability  of  the  horns  of  deer,  as  notably 
in  the  wild  reindeer — the  occasional  presence  of  horns  in  the  fe- 
male Antilope  bezoartica,  and  their  frequent  absence  in  the  female 
of  Antilocapra  americana — the  presence  of  two  tusks  in  some  few 
male  narwhals — the  complete  absence  of  tusks  in  some  female 
walruses — are  all  instances  of  the  extreme  variability  of  secondary 
sexual  characters,  and  of  their  liability  to  differ  in  closely-allied 
forms. 

Although  tusks  and  horns  appear  in  all  cases  to  have  been  pri- 
marily developed  as  sexual  weapons,  they  often  serve  other  pur- 
poses. The  elephant  uses  his  tusks  in  attacking  the  tiger;  ac- 
cording to  Bruce,  he  scores  the  trunks  of  trees  until  they  can  be 
thrown  down  easily,  and  he  likewise  thus  extracts  the  farinaceous 
cores  of  palms;  in  Africa  he  often  uses  one  tusk,  always  the  same, 
to  probe  the  ground  and  thus  ascertain  whether  it  will  bear  his 
weight.  The  common  bull  defends  the  herd  with  his  horns;  and  the 
elk  in  Sweden  has  been  known,  according  to  Lloyd,  to  strike  a  wolf 
dead  with  a  single  blow  of  his  great  horns.  Many  similar  facts 
could  be  given.  One  of  the  most  curious  secondary  uses  to  which 
the  horns  of  an  animal  may  be  occasionally  put,  is  that  observed 
by  Captain  Hutton-^  with  the  wild  goat  (Capra  segagrus)  of  the 
Himalayas,  and  as  it  is  also  said,  with  the  ibex,  namely,  that  when 
the  male  accidentally  falls  from  a  height  he  bends  inwards  his 
head,  and  by  alighting  on  his  massive  horns  breaks  the  shock. 
The  female  cannot  thus  use  her  horns,  which  are  smaller,  but  from 


18  I  have  given  various  experiments  and  other  evidence  proving 
that  this  is  the  case,  in  my  'Variation  of  Animals  and  Plants  under 
Pomestication,'  vol.  ii.,  1868,  pp.  39-47. 

^-  Sir  J.  Emerson  Tennent,  'Ceylon,'  1859,  vol.  ii.  p.  274.  For  Malacca, 
'Journal  of  Indian  Archipelago,'  vol.  iv.  p.  357. 

21  'Calcutta  Journal  of  Nat.  Hist.'  vol.  ii.  1843,  p.  526. 


504  THE  DESCENT  OF  MAN. 

her  more  quiet  disposition  she  does  not  need  this  strange  kind  of 
shield  so  much. 

Each  male  animal  uses  his  weapons  in  his  own  peculiar  fashion. 
The  common  ram  makes  a  charge  and  butts  with  such  force  with 
the  bases  of  his  horns,  that  I  have  seen  a  powerful  man  knocked 
over  like  a  child.  Goats  and  certain  species  of  sheep,  for  in- 
stance the  Ovis  cycloceros  of  Afghanistan,"  rear  on  their  hind 
legs,  and  then  not  only  butt,  but  "make  a  cut  down  and  a  jerk  up. 
"with  the  ribbed  front  of  their  scimitar-shaped  horn,  as  with  a 
"siaber.  When  the  O.  cycloceros  attacked  a  large  domestic  ram, 
"who  was  a  noted  bruiser,  he  conquered  him  by  the  sheer  novelty 
"of  his  mode  of  fighting,  always  closing  at  once  with  his  adver- 
"sary,  and  catching  him  across  the  face  and  nose  with  a  sharp 
"drawing  jerk  of  the  head,  and  then  bounding  out  of  the  way  be- 
"fore  the  blow  could  be  returned."  In  Pembrokeshire  a  male  goat, 
the  master  of  a  flock  which  during  several  generations  had  run 
wild,  was  known  to  have  killed  several  males  in  single  combat; 
this  goat  possessed  enormous  horns,  measuring  thirty-nine  inches 
in  a  straight  line  from  tip  to  tip.  The  common  bull,  as  every  one 
knows,  gores  and  tosses  his  opponent;  but  the  Italian  buffalo  is 
said  never  to  use  his  horns,  he  gives  a  tremendous  blow  with  his 
convex  forehead,  and  then  tramples  on  his  fallen  enemy  with  his 
knees— an  instinct  which  the  common  bull  does  not  possess."-'^ 
Hence  a  dog  who  pins  a  buffalo  by  the  nose  is  immediately  crush- 
ed. We  must,  however,  remember  that  the  Italian  buffalo  has  been 
long  domesticated,  and  it  is  by  no  means  certain  that  the 
wild  parent-form  had  similar  horns.  Mr.  Bartlett  informs  me 
that  when  a  female  Cape  buffalo  (Bubalus  caffer)  was  turned  into 
an  enclosure  with  a  bull  of  the  same  species,  she  attacked  him,  and 
he  in  return  pushed  her  about  with  great  violence.  But  it  was 
manifest  to  Mr.  Bartlett  that,  had  not  the  bull  shown  dignified 
forbearance,  he  could  easily  have  killed  her  by  a  single  lateral 
thrust  with  his  immense  horns.  The  giraffe  uses  his  short  hair- 
covered  horns,  which  are  rather  longer  in  the  male  than  in  the 
female,  in  a  curious  manner;  for,  with  his  long  neck,  he  swings  his 
head  to  either  side,  almost  upside  down,  with  such  force,  that  I 
have  seen  a  hard  plank  deeply  indented  by  a  single  blow. 

With  antelopes  it  is  sometimes  difficult  to  imagine  how  they  can 
possibly  use  their  curiously-shaped  horns;  thus  the  spring-boc 
(Ant.  euchore)  has  rather  short  upright  horns,  with  the  sharp 
points  bent  inwards  almost  at  right  angles,  so  as  to  face  each 


22  Mr.  Blylh,  in  'Land  and  Water,'  March,  1867,  p.  134,  on  the  au- 
thority of  Capt.  Hutton  and  others.  For  the  wild  Pembrokeshire 
goats,  see  the  'Field,'  1869,  p.  150. 

23  M.  E,  M.  Bailly,  'Sur  1' usage  des  Cornes,'  &c.,  'Annal.  des  Sc. 
Nat.'  torn.  ii.  1824,  p.  369. 


MAMMALS— LAW  OF  BATTLE. 


505 


other;  Mr.  Bartlett  does  not  know  how  they  are  used,  but  sug- 
gests that  they  would  inflict  a  fearful  wound  down  each  side  of  the 
face  of  an  antagonist.  The  slightly-curved  horns  of  the  Oryx 
leucoryx  (fig.  63)  are  directed  backwards,  and  are  of  such  length 
that  their  points  reach  beyond  the  middle  of  the  back,  over  which 
they  extend  in  almost  parallel  lines.  Thus  they  seem  singularly 
ill-fitted  for  fighting;  but  Mr.  Bartlett  informs  me  that  when  two 
of  these  animals  prepare  for  battle,  they  kneel  down,  with  their 
heads  between  their  fore  legs,  and  in  this  attitude  the  horns  stand 
nearly  parallel  and  close  to  the  ground,  with  the  points  directed 
forwards  and  a  little  upwards.  The  combatants  then  gradually 
approach  each  other,  and  each  endeavors  to  get  the  upturned 


Fig.  63.    Oryx  leucoryx,  male  (from  the  Knowsley  Menag-erie). 

points  under  the  body  of  the  other;  if  one  succeeds  in  doing  this, 
he  suddenly  springs  up,  throwing  up  his  head  at  the  same  time, 
and  can  thus  wound  or  perhaps  even  transfix  his  antagonist.  Both 
animals  always  kneel  down,  so  as  to  guard  as  far  as  possible 
against  this  maneuver.  It  has  been  recorded  that  one  of  these 
antelopes  has  used  his  horns  with  effect  even  against  a  lion;  yet 
from  being  forced  to  place  his  head  between  the  fore-legs  in  order 
to  bring  the  points  of  the  horns  forward,  he  would  generally  be 
under  a  great  disadvantage  when  attacked  by  any  other  animal. 
It  is,  therefore,  not  probable  that  the  horns  have  been  modified 
into  their  present  great  length  and  peculiar  position,  as  a  protec- 
tion against  beasts  of  prey.  We  can  however  see  that,  as  soon  as 
some  ancient  male  progenitor  of  the  Oryx  acquired  moderately 
long  horns,  directed  a  little  backwards,  he  would  be  compelled,  in 
his  battles  with  rival  males,  to  bend  his  head  somewhat  inwards 
or  downwards,  as  is  now  done  by  certain  stags;  and  it  is  not  im- 
probable that  he  might  have  acquired  the  habit  of  at  first  occa- 


506  THE  DESCENT  OF  MAN. 

sionally  and  afterwards  of  regularly  kneeling  down.  In  this  case 
it  is  almost  certain  that  the  males  which  possessed  the  longest 
horns  would  have  had  a  great  advantage  over  others  with  shorter 
horns;  and  then  the  horns  v/ould  gradually  have  been  rendered 
longer  and  longer,  through  sexual  selection,  until  they  acquired 
their  present  extraordinary  length  and  position. 

With  stags  of  many  kinds  the  branches  of  the  horns  offer  a 
curious  case  of  difficulty;  for  certainly  a  single  straight  point 
would  inflict  a  much  more  serious  wound  than  several  diverging 
ones.  In  Sir  Philip  Egerton's  museum  there  is  a  horn  of  the  red- 
deer  (Cervus  elaphus),  thirty  inches  in  length,  with  "not  fewer 
"than  fifteen  snags  or  branches;"  and  at  Moritzburg  there  is  still 
preserved  a  pair  of  antlers  of  a  red-deer,  shot  in  1699  by  Fred- 
erick I.,  one  of  which  bears  the  astonishing  number  of  thirty- 
three  branches  and  the  other  twenty-seven,  making  altogether 
sixty  branches.  Richardson  figures  a  pair  of  antlers  of  the  wild 
reindeer  with  twenty-nine  points.^*  From  the  manner  in  which 
the  horns  are  branched,  and  more  especially  from  deer  being 
known  occasionally  to  fight  together  by  kicking  with  their  fore- 
feet,-^ M.  Bailly  actually  comes  to  the  conclusion  that  their  horns 
are  more  injurious  than  useful  to  them?  But  this  author  over- 
looks the  pitched  battles  between  rival  males.  As  I  felt  much 
perplexed  about  the  use  or  advantage  of  the  branches,  I  applied 
to  Mr.  McNeill  of  Colonsay,  who  has  long  and  carefully  observed 
the  habits  of  red-deer,  and  he  informs  me  that  he  has  never  seen 
some  of  the  branches  brought  into  use,  but  that  the  brow  antlers, 
from  inclining  downwards,  are  a  great  protection  to  the  forehead, 
and  their  points  are  likewise  used  in  attack.  Sir  Philip  Egerton 
also  informs  me  both  as  to  red-deer  and  fallow-deer  that,  in  fight- 
ing, they  suddenly  dash  together,  and  getting  their  horns  fixed 
against  each  other's  bodies,  a  desperate  struggle  ensues.  When 
one  is  at  last  forced  to  yield  and  turn  round,  the  victor  endeavors 
to  plunge  his  brow  antlers  into  his  defeated  foe.  It  thus  appears 
that  the  upper  branches  are  used  chiefly  or  exclusively  for  pushing 
and  fencing.  Nevertheless  in  some  species  the  upper  branches 
are  used  as  weapons  of  offense;  when  a  man  was  attacked  by  a 
wapiti  deer  (Cervus  canadensis)  in  Judge  Caton's  park  in  Ottawa, 


2*  On  the  horns  of  red-deer,  Owen,  'British  Fossil  Mammals.'  1843, 
p.  478;  Pdchardson  on  the  horns  of  the  reindeer,  'Fauna  Bor.  Ameri- 
cana,' 1829,  p.  240.  I  am  indebted  to  Prof.  Victor  Cams,  for  the  Moritz- 
burg case. 

25  Hon.  J.  D.  Caton  ('Ottawa  Acad,  of  Nat.  Science,'  May,  1868,  p.  9), 
saj's  that  the  American  deer  fight  with  their  fore-feet,  after  "the 
"question  of  superiority  has  been  once  settled  and  acknowledged  in 
"the  herd."  Bailly,  'Sur  I'usage  des  Cornes,'  'Annales  des  Sc.  Nat.' 
torn.  ii.  1824,  p.  371. 


MAMMALS— LAW  OF  BATTLE. 


507 


and  several  men  tried  to  rescue  him,  the  stag  "never  raised  his 
"head  from  the  ground;  in  fact,  he  kept  his  face  almost  flat  on 
"the  ground,  with  his  nose  nearly  between  his  fore  feet,  except 
"when  he  rolled  his  head  to  one  side  to  take  a  new  observation 


Fig.   64.    Strepsiceros   Kudu   (from  Sir  Andrew   Smith's    'Zoology  of 

South  Africa'). 

"preparatory  to  a  plunge."  In  this  position  the  ends  of  the  horns 
were  directed  against  his  adversaries.  "In  rolling  his  head  he 
"necessarily  raised  it  somewhat,  because  his  antlers  were  so  long 
"that  he  could  not  roll  his  head  without  raising  them  on  one  side, 
"while,  on  the  other  side,  they  touched  the  ground."    The  stag 


508  THE  DESCENT  OP  MAN. 

by  this  procedure  gradually  drove  the  party  of  rescuers  backwards, 
to  a  distance  of  150  or  200  feet;  and  the  attacked  man  was  killed.==« 

Although  the  horns  of  stags  are  efficient  weapons,  there  can, 
I  think  be  no  doubt  that  a  single  point  would  have  been  much 
more  dangerous  than  a  branched  antler;  and  Judge  Caton,  who 
has  had  large  experience  with  deer,  fully  concurs  in  this  conclu- 
sion. Nor  do  the  branching  horns,  though  highly  important  as 
a  means  of  defense  against  rival  stags,  appear  perfectly  well 
adapted  for  this  purpose,  as  they  are  liable  to  become  interlocked. 
The  suspicion  has  therefore  crossed  my  mind  that  they  may  serve 
in  part  as  ornaments.  That  the  branched  antlers  of  stags  as  well 
as  the  elegant  lyrated  horns  of  certain  antelopes,  with  their 
graceful  double  curvature,  (fig.  64),  are  ornamental  in  our  eyes, 
no  one  will  dispute.  If  then,  the  horns,  like  the  splendid  accoutre- 
ments of  the  knights  of  old,  add  to  the  noble  appearance  of  stags 
and  antelopes,  they  may  have  been  modified  partly  for  this  pur- 
pose, though  mainly  for  actual  service  in  battle;  but  I  have  no 
evidence  in  favor  of  this  belief. 

An  interesting  case  has  lately  been  published,  from  which  it  ap- 
pears that  the  horns  of  a  deer  in  one  district  in  the  United  States 
are  now  being  modified  through  sexual  and  natural  selection.  A 
writer  in  an  excellent  American  Journal"  says,  that  he  has  hunted 
for  the  last  twenty-one  years  in  the  Adirondacks,  where  the 
Cervus  virginianus  abounds.  About  fourteen  years  ago  he  first 
heard  of  spike-horn  bucks.  These  became  from  year  to  year  more 
common;  about  five  years  ago  he  shot  one,  and  afterwards  an- 
other, and  now  they  are  frequently  killed.  "The  spike-horn  dif- 
"fers  greatly  from  the  common  antler  of  the  C.  virginianus.  It  con- 
"sists  of  a  single  spike,  more  slender  than  the  antler,  and  scarcely 
"half  so  long,  projecting  forward  from  the  brow,  and  terminating 
"in  a  very  sharp  point.  It  gives  a  considerable  advantage  to  its  pos- 
"sessor  over  the  common  buck.  Besides  enabling  him  to  run  more 
"swiftly  through  the  thick  woods  and  underbrush  (every  hunter 
"knows  that  does  and  yearling  bucks  run  much  more  rapidly  than 
"the  large  bucks  when  armed  with  their  cumbrous  antlers),  the 
"spike-horn  is  a  more  effective  weapon  than  the  common  antler. 
"With  this  advantage  the  spike-horn  bucks  are  gaining  upon  the 
"common  bucks,  and  may,  in  time,  entirely  supersede  them  in  the 
"Adirondacks.  Undoubtedly  the  first  spike-horn  buck  was  merely 
"an  accidental  freak  of  nature.  But  his  spike-horns  gave  him  an 
"advantage,  and  enabled  him  to  propagate  his  peculiarity.  His 
"descendants  having  a  like  advantage,  have  propagated  the  pe- 
"culiarity  in  a  constantly  increasing  ratio,  till  they  are  slowly 


2«See  a  most  interesting  account  in  the  Appendix  to  Hon.   J.  D. 
Caton' s  paper,  as  above  quoted. 
"  'The  American  Naturalist,'  Dec.  1869,  p.  552. 


MAMMALS— LAW  OF  BATTLE.  509 

"crowding  the  antlered  deer  from  the  region  they  inhabit."  A 
critic  has  well  objected  to  this  account  by  asking,  why,  if  the 
simple  horns  are  now  so  advantageous,  were  the  branched  antlers 
of  the  parent-form  ever  developed?  To  this  I  can  only  answer 
by  remarking,  that  a  new  mode  of  attack  with  new  weapons  might 
be  a  great  advantage,  as  shown  by  the  case  of  the  Ovis  cycloceros, 
who  thus  conquered  a  domestic  ram  famous  for  his  fighting  power. 
Though  the  branched  antlers  of  a  stag  are  well  adapted  for  fight- 
ing with  his  rivals,  and  though  it  might  be  an  advantage  to  the 
prong-horned  variety  slowly  to  acquire  long  and  branched  horns, 
if  he  had  to  fight  only  with  others  of  the  same  kind,  yet  it  by  no 
means  follows  that  branched  horns  would  be  the  best  fitted  for 
conquering  a  foe  differently  armed.  In  the  foregoing  case  of  the 
Oryx  leucoryx,  it  is  almost  certain  that  the  victory  would  rest 
with  an  antelope  having  short  horns,  and  who  therefore  did  not 
need  to  kneel  down,  though  an  oryx  might  profit  by  having  still 
longer  horns,  if  he  fought  only  with  his  proper  rivals. 

Male  quadrupeds,  which  are  furnished  with  tusks,  use  them  in 
various  ways,  as  in  the  case  of  horns.  The  boar  strikes  laterally 
and  upwards;  the  musk-deer  downwards  with  serious  effect.^^  The 
walrus,  though  having  so  short  a  neck  and  so  unwieldy  a  body, 
"can  strike  either  upwards,  or  downwards,  or  sideways,  with 
"equal  dexterity."^  I  was  informed  by  the  late  Dr.  Falconer,  that 
the  Indian  elephant  fights  in  a  different  manner  according  to  the 
position  and  curvature  of  his  tusks.  When  they  are  directed  for- 
wards and  upwards  he  is  able  to  fling  a  tiger  to  a  great  distance — 
it  is  said  to  even  thirty  feet;  when  they  are  short  and  turned  down- 
wards he  endeavors  suddenly  to  pin  the  tiger  to  the  ground  and, 
in  consequence,  is  dangerous  to  the  rider,  who  is  liable  to  be 
jerked  off  the  howdah.^** 

Very  few  male  quadrupeds  possess  weapons  of  two  distinct 
kinds  specially  adapted  for  fighting  with  rival  males.  The  male 
muntjac-deer  (Cervulus),  however,  offers  an  exception,  as  he  is 
provided  with  horns  and  exserted  canine  teeth.  But  we  may  infer 
from  what  follows  that  one  form  of  weapon  has  often  been  re- 
placed in  the  course  of  ages  by  another.  With  ruminants  the  de- 
velopment of  horns  generally  stands  in  an  inverse  relation  with 
that  of  even  moderately  developed  canine  teeth.  Thus  camels, 
guanacoes,  chevrotains,  and  musk-deer,  are  hornless,  and  they 
have  efficient  canines;  these  teeth  being  "always  of  smaller  size 
"in  the  females  than  in  the  males."  The  Camelidae  have,  in  ad- 
dition to  their  true  canines,  a  pair  of  canine-shaped  incisors  in 


28  Pallas,  'Spicilegia  Zoologica,*  fasc.  xiii.  1779,  p.  18. 

29  Lament,  'Seasons  with  the  Sea-Horses,'  1861,  p.  141. 

30  See,  also.  Corse  ('Philosoph.  Transact.'  1799,  p.  212)  on  the  manner 
in  which  the  short-tusked  Mooknah  variety  attacks  other  elephants. 


510  THE  DESCENT  OF  MAN. 

their  upper  jaws.'^  Male  deer  and  antelopes,  on  the  other  hand, 
possess  horns,  and  they  rarely  have  canine  teeth;  and  these,  when 
present,  are  always  of  small  size,  so  that  it  is  doubtful  whether 
they  are  of  any  service  in  their  battles.  In  Antilope  montana 
they  exist  only  as  rudiments  in  the  young  male,  disappearing  as 
he  grows  old;  and  they  are  absent  in  the  female  at  all  ages;  but 
the  females  of  certain  other  antelopes  and  of  certain  deer  have 
been  known  occasionally  to  exhibit  rudiments  of  these  teeth,^^ 
Stallions  have  small  canine  teeth,  which  are  either  quite  absent  or 
rudimentary  in  the  mare;  but  they  do  not  appear  to  be  used  in 
fighting,  for  stallions  bite  with  their  incisors,  and  do  not  open 
their  mouths  wide  lil^e  camels  and  guanacoes.  Whenever  the 
adult  male  possesses  canines,  now  inefficient,  whilst  the  female  has 
either  none  or  mere  rudiments,  we  may  conclude  that  the  early 
male  progenitor  of  the  species  was  provided  with  efficient  canines, 
which  have  been  partially  transferred  to  the  females.  The  re- 
duction of  these  teeth  in  the  males  seems  to  have  followed  from 
some  change  in  their  manner  of  fighting,  often  (but  not  in  the 
horse)  caused  by  the  development  of  new  weapons. 

Tusks  and  horns  are  manifestly  of  high  importance  to  their 
possessors,  for  their  development  consumes  much  organized  mat- 
ter. A  single  tusk  of  the  Asiatic  elephant — one  of  the  extinct 
woolly  species — and  of  the  African  elephant,  have  been  known  to 
weigh  respectively  150,  160,  and  180  pounds;  and  even  greater 
weights  have  been  given  by  some  authors.^^  With  dfeer,  in  which 
the  horns  are  periodically  renewed,  the  drain  on  the  constitution 
must  be  greater;  the  horns,  for  instance,  of  the  moose  weigh  from 
fifty  to  sixty  pounds,  and  those  of  the  extinct  Irish  elk  from  sixty 
to  seventy  pounds — the  skull  of  the  latter  weighing  on  an  average 
only  five  pounds  and  a  quarter.  Although  the  horns  are  not  pe- 
riodically renewed  in  sheep,  yet  their  development,  in  the  opin- 
ion of  many  agriculturists,  entails  a  sensible  loss  to  the  breeder. 
Stags,  moreover,  in  escaping  from  beasts  of  prey  are  loaded  with 
an  additional  weight  for  the  race,  and  are  greatly  retarded  in 
passing  through  a  woody  country.  The  moose,  for  instance,  with 
horns  extending  five  and  a  half  feet  from  tip  to  tip,  although  so 


31  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  349. 

32  See  Ruppell  (in  'Proc.  Zoolog.  Soc'  Jan.  12,  1836,  p.  3)  on  the  canines 
in  deer  and  antelopes,  with  a  note  by  Mr.  Martin  on  a  female  Ameri- 
can deer.  See,  also.  Falconer  ('Palaeont.  Memoirs  and  Notes,'  vol.  i. 
1868,  p.  576)  on  canines  in  an  adult  female  deer.  In  old  males  of  the 
musk-deer  the  canines  (Pallas,  'Spic.  Zoolog.'  fasc.  xiii.  1779,  p.  18) 
sometimes  grow  to  the  length  of  three  inches,  whilst  in  old  females 
a  rudiment  projects  scarcely  half  an  inch  above  the  gums. 

33  Emerson  Tennent,  'Ceylon,'  1859,  vol.  ii.  p.  275;  Owen,  'British  Fos- 
sil Mammals,'  1846,  p.  245. 


MAMMALS— GREATER    SIZE.  OF    MALE.  511 

skillful  in  their  use  that  he  will  not  touch  or  break  a  twig  when 
walking  quietly,  cannot  act  so  dexterously  whilst  rushing  away 
from  a  pack  of  wolves.  "During  his  progress  he  holds  his  nose 
"up  so  as  to  lay  the  horns  horizontally  back;  and  in  this  atti- 
"tude  cannot  see  the  ground  distinctly. "=^*  The  tips  of  the  horns  of 
the  great  Irish  elk  were  actually  eight  feet  apart!  Whilst  the 
horns  are  covered  with  velvet,  which  lasts  with  the  red-deer  for 
about  twelve  weeks,  they  are  extremely  sensitive  to  a  blow;  so 
that  in  Germany  the  stags  at  this  time  somewhat  change  their 
habits,  and  avoiding  dense  forests,  frequent  young  woods  and 
low  thickets. ^^  These  facts  remind  us  that  male  birds  have  ac- 
quired ornamental  plumes  at  the  cost  of  retarded  flight,  and  other 
ornaments  at  the  cost  of  some  loss  of  power  in  their  battles  with 
rival  males. 

With  mammals,  when,  as  is  often  the  case,  the  sexes  differ  in 
size,  the  males  are  almost  always  larger  and  stronger.  I  am  in- 
formed by  Mr.  Gould,  that  this  holds  good  in  a  marked  manner 
with  the  marsupials  of  Australia,  the  males  of  which  appear  to 
continue  growing  until  an  unusually  late  age.  But  the  most  ex- 
traordinary case  is  that  of  one  of  the  seals  (Callorhinus  ursinus), 
a  full-grown  female  weighing  less  than  one-sixth  of  a  full-grown 
male.^^  Dr.  Gill  remarks  that  it  is  with  the  polygamous  seals,  the 
males  of  which  are  well  known  to  fight  savagely  together,  that 
the  sexes  differ  much  in  size;  the  monogamous  species  differing 
but  little.  Whales  also  afford  evidence  of  the  relation  existing 
between  the  pugnacity  of  the  males  and  their  large  size  compared 
with  that  of  the  female;  the  males  of  the  right- whales  do  not 
fight  together,  and  they  are  not  larger,  but  rather  smaller,  than 
their  females;  on  the  other  hand,  male  sperm-whales  fight  much 
together,  and  their  bodies  are  "often  found  scarred  with  the  im- 
"print  of  their  rival's  teeth,"  and  they  are  double  the  size  of  the 
females.  The  greater  strength  of  the  male,  as  Hunter  long  ago 
remarked,"'  is  invariably  displayed  in  those  parts  of  the  body 
which  are  brought  into  action  in  fighting  with  rival  males — for 
instance,  in  the  massive  neck  of  the  bull.    Male  quadrupeds  are 


8*  Richardson,  'Fauna  Bor.  Americana,'  on  the  moose,  Alces  pal- 
mata,  pp.  236,  237;  on  the  expanse  of  the  horns,  'Land  and  Water.' 
1869,  p.  143.  See,  also,  Owen,  'British  Fossil  Mammals,'  on  the  Irish 
elk,  pp.  447,  455. 

35  'Forest  Creatures,'  by  C.  Boner,  1861,  p.  60. 

36  See  the  very  interesting-  paper  by  Mr.  J.  A.  Allen  in  'Bull.  Mus. 
Comp.  Zoolog.  of  Cambridge,  United  States,'  vol.  ii.  No.  1,  p.  82.  The 
weights  were  ascertained  by  a  careful  observer,  Capt.  Bryant.  Dr^ 
Gill  in  'The  American  Naturalist,'  Jan.  1871,  Prof.  Slialer  on  the  rela- 
tive size  of  the  sexes  of  whales,  'American  Naturalist,'  Jan.  1873. 

^''  'Animal  Economy,'  p.  45. 


512  THE  DESCENT  OF  MAN. 

also  more  courageous  and  pugnacious  than  the  females.  There 
can  be  little  doubt  that  these  characters  have  been  gained,  partly 
through  sexual  selection,  owing  to  a  long  series  of  victories  by 
the  stronger  and  more  courageous  males  over  the  weaker,  and 
partly  through  the  inherited  effects  of  use.  It  is  probable  that 
the  successive  variations  in  strength,  size,  and  courage,  whether 
due  to  mere  variability  or  to  the  effects  of  use,  by  the  accumula- 
tion of  which  male  quadrupeds  have  accLuired  these  characteristic 
qualities,  occurred  rather  late  in  life,  and  were  consequently  to  a 
large  extent  limited  in  their  transmission  to  the  same  sex. 

From  these  considerations  I  was  anxious  to  obtain  information 
as  to  the  Scotch  deerhound,  the  sexes  of  which  differ  more  in 
size  than  those  of  any  other  breed  (though  bloodhounds  differ 
considerably),  or  than  in  any  wild  canine  species  known  to  me. 
Accordingly,  I  applied  to  Mr.  Cupples,  well-known  for  his  suc- 
cess with  this  breed,  who  has  weighed  and  measured  many  of 
his  own  dogs,  and  who  has  with  great  kindness  collected  for  me 
the  following  facts  from  various  sources.  Fine  male  dogs,  meas- 
ured at  the  shoulder,  range  from  28  inches,  which  is  low,  to  33, 
or  even  34  inches  in  height;  and  in  weight  from  80  pounds,  which 
is  light,  to  120  pounds,  or  even  more.  The  females  range  in 
height  from  23  to  27,  or  even  to  28  inches;  and  in  weight  from 
50  to  70,  or  even  80  pounds.^^  Mr.  Cupples  concludes  that  from 
95  to  100  pounds  for  the  male,  and  70  for  the  female,  would  be  a 
safe  average;  but  there  is  reason  to  believe  that  formerly  both 
sexes  attained  a  greater  weight.  Mr.  Cupples  has  weighed  pup- 
pies when  a  fortnight  old;  in  one  litter  the  average  weight  of 
four  males  exceeded  that  of  two  females  by  six  and  a  half  ounces; 
in  another  litter  the  average  weight  of  four  males  exceeded  that 
of  one  female  by  less  than  one  ounce;  the  same  males  when 
three  weeks  old,  exceeded  the  female  by  seven  and  a  half  ounces, 
and  at  the  age  of  six  weeks  by  nearly  fourteen  ounces.  Mr. 
Wright  of  Yeldersley  House,  in  a  letter  to  Mr.  Cupples,  says:  "1 
"have  taken  notes  on  the  sizes  and  weights  of  puppies  of  many 
"litters,  and  as  far  as  my  experience  goes,  dog-puppies  as  a  rule 
"differ  very  little  from  bitches  till  they  arrive  at  about  five  or  six 
"months  old;  and  then  the  dogs  begin  to  increase,  gaining  upon 
"the  bitches  both  in  weight  and  size.  At  birth,  and  for  several 
"weeks  afterwards,  a  bitch-puppy  will  occasionally  be  larger  than 
"any  of  the  dogs,  but  they  are  invariably  beaten  by  them  later." 


38  See,  also,  Richardson's  'Manual  on  the  Dog,'  p.  59.  Much  valuable 
information  on  the  Scottish  deer-hound  is  given  by  Mr.  McNeill,  who 
first  called  attention  to  the  inequality  in  size  between  the  sexes,  in 
Scrope's  'Art  of  Deer  Stalking.'  I  hope  that  Mr.  Cupples  will  keep 
to  his  intention  of  publishing  a  full  account  and  history  of  this 
famous  breed. 


MAMMALS— GREATER  SIZE   OF  MALE. 


513 


Mr.  McNeill,  of  Colonsay,  concludes  that  "the  males  do  not  at- 
"tain  their  full  growth  till  over  two  years  old,  though  the  fe- 
"males  attain  it  sooner."  According  to  Mr.  Cupples'  experience, 
male  dogs  go  on  growing  in  stature  till  they  are  from  twelve  to 
eighteen  months  old,  and  in  weight  till  from  eighteen  to  twenty- 
four  months  old;  whilst  the  females  cease  increasing  in  stature 
at  the  age  of  from  nine  to  fourteen  or  fifteen  months,  and  in  weight 
at  the  age  of  from  twelve  to  fifteen  months.  From  these  various 
statements  it  is  clear  that  the  full  difference  in  size  between  the 
male  and  female  Scotch  deerhound  is  not  acquired  until  rather 
late  in  life.  The  males  almost  exclusively  are  used  for  coursing, 
for,  as  Mr.  McNeill  informs  me,  the  females  have  not  sufiicient 
strength  and  weight  to  pull  down  a  full-grown  deer.  From  the 
names  used  in  old  legends,  it  appears,  as  I  hear  from  Mr.  Cup- 
ples, that,  at  a  very  ancient  period,  the  males  were  the  most  cele- 
brated, the  females  being 
mentioned  only  as  the  moth- 
ers of  famous  dogs.  Hence, 
during  many  generations,  it 
is  the  male  which  has  been 
chiefly  tested  for  strength, 
size,  speed,  and  courage,  and 
the  best  will  have  been  bred 
from.  As,  however,  the  males 
do  not  attain  their  full  dimen- 
sions until  rather  late  in  life, 
they  will  have  tended,  in  ac- 
cordance with  the  law  often 
indicated,  to  transmit  their 
characters  to  their  male  off- 
spring alone;  and  thus  the 
great  inequality  in  size  be- 
tween the  sexes  of  the  Scotch  deerhounds  may  probably  be  ac- 
counted for. 

The  males  of  some  few  quadrupeds  possess  organs  or  parts 
developed  solely  as  a  means  of  defense  against  the  attacks  of 
other  males.  Some  kinds  of  deer  use,  as  we  have  seen,  the  upp^^^ 
branches  of  their  horns  chiefly  or  exclusively  for  defending  them- 
selves; and  the  Oryx  antelope,  as  I  am  informed  by  Mr.  Bartlett. 
fences  most  skillfully  with  his  long,  gently  curved  horns;  but 
these  are  likewise  used  as  organs  of  offense.  The  same  observer 
remarks  that  rhinoceroses  in  fighting,  parry  each  other's  sidelong 
blows  with  their  horns,  which  clatter  loudly  together,  as  do  the 
tusks  of  boars.  Although  wild  boars  fight  desperately,  they  sel- 
dom, according  to  Brehm,  receive  fatal  wounds,  as  the  blows  fall 
on  each  other's  tusks,  or  on  the  layer  of  gristly  skin  covering  the 
shoulder,  called  by  the  German  hunters,  the  shield;  and  here 
34  '  . 


•r>SgsC_>5i 


Fig.  85.    Head  of  Common  wild  boar, 
in  prime  of  life  (from  Brehm). 


514 


THE  DESCENT  OF  MAN. 


we  have  a  part,  specially  modified  for  defense.  With  boars  in 
the  prime  of  life  (see  fig.  65)  the  tusks  in  the  lower  jaw  are  used 
for  fighting,  but  they  become  in  old  age,  as  Brehm  states,  so  much 
curved  inwards  and  upwards  over  the  snout,  that  they  can  no 
longer  be  used  in  this  way.  They  may,  however,  still  serve,  and 
even  more  effectively,  as  a  means  of  defense.    In  compensation 


Fig.  66.    Skull  of  the  Babirusa  Pig  (from  Wallace's  'Malay  Archi- 
pelago'). 

for  the  loss  of  the  lower  tusks  as  weapons  of  offense,  those  in 
the  upper  jaw,  which  always  project  a  little  laterally,  increase  in 
old  age  so  much  in  length  and  curve  so  much  upwards,  that  they 
can  be  used  for  attack.  Nevertheless,  an  old  boar  is  not  so  dan- 
gerous to  man  as  one  at  the  age  of  six  or  seven  years.^^ 

In  the  full-grown  male  Babirusa  pig  of  Celebes  (fig.  66),  the 
lower  tusks  are  formidable  weapons,  like  those  of  the  European 
boar  in  the  prime  of  life,  whilst  the  upper  tusks  are  so  long 
and  have  their  points  so  much  curved  inwards,  sometimes  even 
touching  the  forehead,  that  they  are  utterly  useless  as  weapons 
of  attack.    They  more  nearly  resemble  horns  than  teeth,  and 


89  Brehm,  'Thierleben,'  B.  ii.  s.  729-732. 


MAMMALS— MEANS    OF    DEFENSE. 


516 


are  so  manifestly  useless  as  teeth,  that  the  animal  was  formerly- 
supposed  to  rest  his  head  by  hooking  them  on  to  a  branch! 
Their  convex  surfaces,  however,  if  the  head  were  held  a  little 
laterally,  would  serve  as  an  excellent  guard;  and  hence,  perhaps, 
it  is  that  in  old  animals  they  "are  generally  broken  off,  as  if  by 
"fighting."*'  Here,  then,  we  have  the  curious  case  of  the  upper 
tusks  of  the  Babirusa  regularly  assuming  during  the  prime  of 
life,  a  structure  which  apparently  renders  them  fitted  only  for 
defense;  whilst  in  the  European  boar  the  lower  tusks  assume  in 
a  less  degree  and  only  during  old  age,  nearly  the  same  form,  and 
then  serve  in  like  manner  solelj'-  for  defense. 


Pig.  67.  Head  of  female  Ethiopian  wart-hog,  from  'Proc.  Zool.  Soc' 
1869,  showing  the  same  characters  as  the  male,  though  on  a  re- 
duced scale. 

N.  B.  When  the  engraving-  was  first  made,  I  was  under  the  impression 
that  it  represented  the  male. 

In  the  wart-hog  (Phacochoerus  sethiopicus,  fig.  67)  the  tusks  in 
the  upper  jaw  of  the  male  curve  upwards  during  the  prime  of 
life  and  from  being  pointed  serve  as  formidable  weapons.  The 
tusks  in  the  lower  jaw  are  sharper  than  those  in  the  upper,  but 
from  their  shortness  it  seems  hardly  possible  that  they  can  be 
used  as  weapons  of  attack.  They  must,  however,  greatly  strength- 
en those  in  the  upper  jaw,  from  being  ground  so  as  to  fit  closely 
against  their  bases.  Neither  the  upper  nor  the  lower  tusks  ap- 
pear to  have  been  specially  modified  to  act  as  guards,  though 
no  doubt  they  are  to  a  certain  extent  used  for  this  purpose. 


^  See  Mr.  Wallace's  interesting  account  of  this  animal,  'The  Malay 
Archipelago,'  1869,  vol.  i.  p.  435. 


516  I'HE  DESCENT  OF  MAN. 

But  the  wart-hog  is  not  destitute  of  other  special  means  of  pro- 
tection, for  it  has,  on  each  side  of  the  face,  beneath  the  eyes,  a 
rather  stiff,  yet  flexible,  cartilaginous,  oblong  pad  (fig.  67),  which 
projects  two  or  three  inches  outwards;  and  it  appeared  to  Mr. 
Bartlett  and  myself,  when  viewing  the  living  animal,  that  these 
pads,  when  struck  from  beneath  by  the  tusks  of  an  opponent, 
would  be  turned  upwards,  and  would  thus  admirably  protect  the 
somewhat  prominent  eyes.  I  may  add,  on  the  authority  of  Mr. 
Bartlett  that  these  boars  when  fighting  stand  directly  face  to 
face. 

Lastly,  the  African  river-hog  (Potomochoerus  penicillatus)  has 
a  hard  cartilaginous  knob  on  each  side  of  the  face  beneath  the 
eyes,  which  answers  to  the  flexible  pad  of  the  wart-hog;  it  has 
also  two  bony  prominences  on  the  upper  jaw  above  the  nostrils. 
A  boar  of  this  species  in  the  Zoological  Gardens  recently  broke 
into  the  cage  of  the  wart-hog.  They  fought  all  night  long,  and 
were  found  in  the  morning  much  exhausted,  but  not  seriously 
wounded.  It  is  a  significant  fact,  as  showing  the  purpose  of  the 
above-described  projections  and  excrescences,  that  these  were 
covered  with  blood,  and  were  scored  and  abraded  in  an  extraor- 
dinary manner. 

Although  the  males  of  so  many  members  of  the  pig  family  are 
provided  with  weapons,  and  as  we  have  just  seen  with  means  of 
defense,  these  weapons  seem  to  have  been  acquired  within  a 
rather  late  geological  period.  Dr.  Forsyth  Major  specifies*^  sev- 
eral miocene  species,  in  none  of  which  do  the  tusks  appear  to 
have  been  largely  developed  in  the  males;  and  Prof.  Riitimeyer 
was  formerly  struck  with  this  same  fact. 

The  mane  of  the  lion  forms  a  good  defense  against  the  attacks 
of  rival  lions,  the  one  danger  to  which  he  is  liable;  for  the 
males,  as  Sir  A.  Smith  informs  me,  engage  in  terrible  battles, 
and  a  young  lion  dares  not  approach  an  old  one.  In  1857  a 
tiger  at  Bromwich  broke  into  the  cage  of  a  lion  and  a  fearful 
scene  ensued:  "the  lion's  mane  saved  his  neck  and  head  from 
"being  much  injured,  but  the  tiger  at  last  succeeded  in  ripping 
"up  his  belly,  and  in  a  few  minutes  he  was  dead."^  The  broad 
ruff  round  the  throat  and  chin  of  the  Canadian  lynx  (Felis 
canadensis)  is  much  longer  in  the  male  than  in  the  female;  but 
whether  it  serves  as  a  defense  I  do  not  know.  Male  seals  are 
well  known  to  fight  desperately  together,  and  the  males  of  cer- 
tain kinds  (Otaria  jubata)*^  have  great  manes,  whilst  the  females 
have  small  ones  or  none.    The  male  baboon  of  the  Cape  of  Good 

«  'Atti  della  Soc.  Italiana  di  Sc.  Nat.'  1873,  vol.  xv.  fasc.  iv. 
*2  'The  Times,'   Nov.  10th,  1857.    In  regard  to  the  Canada  lynx,   see 
Audubon  and  Bachman,  'Quadrupeds  of  N.  America,'  1846,  p.  39. 
"Dr.  Murie,  on  Otaria,   'Proc.  Zoolog.  Soc'  1889,  p.  109.    Mr.  J.  A. 


MAMMALS— CHOICE    IN    PAIRING.  517 

Hope  (Cynocephalus  porcarius)  has  a  much  longer  mane  and 
larger  canine  teeth  than  the  female;  and  the  mane  prohably 
serves  as  a  protection,  for  on  asking  the  keepers  in  the  Zoo- 
logical Gardens,  without  giving  them  any  clue  to  my'  object, 
whether  any  of  the  monkeys  especially  attacked  each  other  by 
the  nape  of  the  neck,  I  was  answered  that  this  was  not  the  case, 
except  with  the  above  baboon.  In  the  Hamadryas  baboon,  Ehren- 
berg  compares  the  mane  of  the  adult  male  to  that  of  a  young 
lion,  whilst  in  the  young  of  both  sexes  and  in  the  female  the 
mane  is  almost  absent. 

-  It  appeared  to  me  probable  that  the  immense  woolly  mane  of 
the  male  American  bison,  which  reaches  almost  to  the  ground, 
and  is  much  more  developed  in  the  males  than  in  the  females, 
served  as  a  protection  to  them  in  their  terrible  battles;  but  an 
experienced  hunter  told  Judge  Caton  that  he  had  never  observed 
anything  which  favored  this  belief.  The  stallion  has  a  thicker 
and  fuller  mane  than  the  mare;  and  I  have  made  particular 
inquiries  of  two  great  trainers  and  breeders,  who  have  had  charge 
of  many  entire  horses,  and  am  assured  that  they  "invariably 
"endeavor  to  seize  one  another  by  the  neck."  It  does  not,  how- 
ever, follow  from  the  foregoing  statements,  that  when  the  hair 
on  the  neck  serves  as  a  defense,  that  it  was  originally  developed 
for  this  purpose,  though  this  is  probable  in  some  cases,  as  in  that 
of  the  lion.  I  am  informed  by  Mr.  McNeill  that  the  long  hairs 
on  the  throat  of  the  stag  (Cervus  elaphus)  serve  as  a  great  pro- 
tection to  him  when  hunted,  for  the  dogs  generally  endeavor  to 
seize  him  by  the  throat;  but  it  is  not  probable  that  these  hairs 
were  specially  developed  for  this  purpose;  otherwise  the  young 
and  the  females  would  have  been  equally  protected. 

Choice  in  Pairing  by  either  Sex  of  Quadrupeds.  — ^Before  de- 
scribing in  the  next  chapter,  the  differences  between  the  sexes  in 
voice,  odors  emitted,  and  ornaments,  it  will  be  convenient  here 
to  consider  whether  the  sexes  exert  any  choice  in  their  unions. 
Does  the  female  prefer  any  particular  male,  either  before  or 
after  the  males  may  have  fought  together  for  supremacy;  or 
does  the  male,  when  not  a  poiygamist,  select  any  particular  fe- 
male? The  general  impression  amongst  breeders  seems  to  be 
that  the  male  accepts  any  female;  and  this  owing  to  his  eager- 
ness, is,  in  most  cases,  probably  the  truth.  Whether  the  female 
as  a  general  rule  indifferently  accepts  any  male  is  much  more 
doubtful.  In  the  fourteenth  chapter,  on  Birds,  a  considerable 
body  of  direct  and  indirect  evidence  was  advanced,  showing  that 

Allen,  in   the  paper  above  quoted   (p.  75),   doubts   whether  the  hair, 
which  is  longer  on  the  neck  in  the  male  than  in  the  female,  deserves 
to  be  called  a  mane. 
34 


518  THE  DESCENT  OF  MAN. 

the  female  selects  her  partner;  and  it  would  be  a  strange  anom- 
aly if  female  quadrupeds,  which  stand  higher  in  the  scale  and 
have  higher  mental  powers,  did  not  generally,  or  at  least  often, 
exert  some  choice.  The  female  could  in  most  cases  escape,  if 
wooed  by  a  male  that  did  not  please  or  excite  her;  and  when  pur- 
sued by  several  males,  as  commonly  occurs,  she  would  often  have 
the  opportunity,  whilst  they  were  fighting  together,  of  escaping 
with  some  one  male,  or  at  least  of  temporarily  pairing  with  him. 
This  latter  contingency  has  often  been  observed  in  Scotland 
with  female  red-deer,  as  I  am  informed  by  Sir  Philip  Egerton 
and  others.** 

It  is  scarcely  possible  that  much  should  be  known  about  fe- 
male quadrupeds  in  a  state  of  nature  making  any  choice  in  their 
marriage  unions.  The  following  curious  details  on  the  courtship 
of  one  of  the  eared  seals  (Callorhinus  ursinus)  are  given*^  on 
the  authority  of  Capt.  Bryant,  who  had  ample  opportunities  for 
observation.  He  says,  "Many  of  the  females  on  their  arrival  at 
"the  island  where  they  breed  appear  desirous  of  returning  to 
"some  particular  male,  and  frequently  climb  the  outlying  rocks 
"to  overlook  the  rookeries,  calling  out  and  listening  as  if  for  a 
"familiar  voice.  Then  changing  to  another  place  they  do  the 
"same  again.  ...  As  soon  as  a  female  reaches  the  shore,  the 
"nearest  male  goes  down  to  meet  her,  making  meanwhile  a  noise 
"like  the  clucking  of  a  hen  to  her  chickens.  He  bows  to  her  and 
"coaxes  her  until  he  gets  between  her  and  the  water  so  that  she 
"cannot  escape  him.  Then  his  •  manner  changes,  and  with  a 
"harsh  growl  he  drives  her  to  a  place  in  his  harem.  This  con- 
"tinues  until  the  lower  row  of  harems  is  nearly  full.  Then  the 
"males  higher  up  select  the  time  when  their  more  fortunate 
"neighbors  are  off  their  guard  to  steal  their  wives.  This  they 
"do  by  taking  them  in  their  mouths  and  lifting  them  over  the 
"heads  of  the  other  females,  and  carefully  placing  them  in  their 
*'own  harem,  carrying  them  as  cats  do  their  kittens.  Those  still 
"higher  up  pursue  the  same  method  until  the  whole  space  is  occu- 
"pied.  Frequently  a  struggle  ensues  between  two  males  for  the 
"possession  of  the  same  female,  and  both  seizing  her  at  once 
"pull  her  in  two  or  terribly  lacerate  her  with  their  teeth.  "When 
"the  space  is  all  filled,  the  old  male  walks  around  complacently 
"reviewing  his  family,  scolding  those  who  crowd  or  disturb  the 

**  Mr.  Boner,  in  his  excellent  description  of  the  habits  of  the  red- 
deer  in  Germany  ('Forest  Creatures,'  1861,  p.  81)  says,  "while  the  stag 
"is  defending  his  rights  against  one  intruder,  another  invades  the 
"sanctuary  of  his  harem,  and  carries  off  trophy  after  trophy,"  Ex- 
actly the  same  thing  occurs  with  seals,  see  Mr.  J.  A.  Allen,  ibid.  p.  100. 

45  Mr.  J.  A.  Allen  in  'Bull.  Mus.  Comp.  Zoolog.  of  Cambridge,  United 
States,'  vol.  ii.  No.  1,  p.  99. 


MAMMADS-CHOICE    IN    PAIEING.  ol9 

"others,  and  fiercely  driving  off  all  intruders.  This  surveillance 
"always  keeps  him  actively  occupied." 

As  so  little  is  known  about  the  courtship  of  anima/ls  in  a  state 
of  nature,  I  have  endeavored  to  discover  how  far  our  domesti- 
cated quadrupeds  evince  any  choice  in  their  unions.  Dogs  offer 
the  best  opportunity  for  observation,  as  they  are  carefully  at- 
tended to  and  well  understood.  Many  breeders  have  expressed 
a  strong  opinion  on  this  head.  Thus,  Mr.  Mayhew  remarks, 
"The  females  are  able  to  bestow  their  affections;  and  tender 
"recollections  are  as  potent  over  them  as  they  are  known  to  be 
"in  other  cases,  where  higher  animals  are  concerned.  Bitches 
"are  not  always  prudent  in  their  loves,  but  are  apt  to  fiing 
"themselves  away  on  curs  of  low  degree.  If  reared  with  a  com- 
"panion  of  vulgar  appearance,  there  often  springs  up  between 
"the  pair  a  devotion  which  no  time  'can  afterwards  subdue.  The 
"passion,  for  such  it  really  is,  becomes  of  a  more  than  romantic 
"endurance."  Mr.  Mayhew,  who  attended  chiefly  to  the  smaller 
breeds,  is  convinced  that  the  females  are  strongly  attracted  by 
males  of  a  large  size.^^  The  well-known  veterinary  Blaine  states*^ 
that  his  own  female  pug  became  so  attached  to  a  spaniel,  and  a 
female  setter  to  a  cur,  that  in  neither  case  would  they  pair  with 
a  dog  of  their  own  breed  until  several  weeks  had  elapsed.  Two 
similar  and  trustworthy  accounts  have  been  given  me  in  regard 
to  a  female  retriever  and  a  spaniel,  both  of  which  became  enam- 
ored with  terrier-dogs. 

Mr.  Cupples  informs  me  that  he  can  personally  vouch  for  the 
accuracy  of  the  following  more  remarkable  case,  in  which  a 
valuable  and  wonderfully  intelligent  female  terrier  loved  a  re- 
triever belonging  to  a  neighbor  to  such  a  degree,  that  she  had 
often  to  be  dragged  away  from  him.  After  their  permanent  sep- 
aration, although  repeatedly  showing  milk  in  her  teats,  she  would 
never  acknowledge  the  courtship  of  any  other  dog,  and  to  the 
regret  of  her  owner  never  bore  puppies.  Mr,  Cupples  also  states, 
that  in  1868,  a  female  deerhound  in  his  kennel  thrice  produced 
puppies,  and  on  each  occasion  showed  a  marked  preference  for 
one  of  the  largest  and  handsomest,  but  not  the  most  eager,  of 
four  deerhounds  living  with  her,  all  in  the  prime  of  life.  Mr. 
Cupples  has  observed  that  the  female  generally  favors  a  dog  whom 
she  has  associated  with  and  knows;  her  shyness  and  timidity 
at  first  incline  her  against  a  strange  dog.  The  male,  on  the 
contrary,  seems  rather  inclined  towards  strange  females.  It  ap- 
pears to  be  rare  when  the  male  refuses  any  particular  female, 
but  Mr.  Wright,  of  Yeldersley  House,  a  great  breeder  of  dogs, 

*6  'Dogs:  their  Management,'  by  E.  Mayhew,  M.  R.  C.  V.  S.,  2nd  edit. 
1864,  pp.  187-192. 

*f  Quoted  by  Alex.  Walker  'On  Intermaxriage,'  183*.  p.  276,  see,  alsQ. 
p.  244. 


520  THE  DESCENT  OF  MAN. 

informs  me  that  he  has  known  some  instances;  he  cites  the  case 
of  one  of  his  own  deerhounds,  who  would  not  take  any  notice 
of  a  particular  female  mastiff,  so  that  another  deerhound  had  to  be 
employed.  It  would  be  superfluous  to  give,  as  I  could,  other 
instances,  and  I  will  only  add  that  Mr.  Barr,  who  has  carefully 
bred  many  bloodhounds,  states  that  in  almost  every  instance 
particular  individuals  of  opposite  sexes  show  a  decided  prefer- 
ence for  each  other.  Finally,  Mr.  Cupples,  after  attending  to 
this  subject  for  another  year,  has  written  to  me,  "I  have  had 
"full  confirmation  of  my  former  statement,  that  dogs  in  breeding 
"form  decided  preferences  for  each  other,  being  often  influenced 
"by  size,  bright  color,  and  individual  characters,  as  well  as  by 
"the  degree  of  their  previous  familiarity." 

In  regard  to  horses,  Mr.  Blenkiron,  the  greatest  breeder  of 
race-horses  in  the  world,  informs  me  that  stallions  are  so  fre- 
quently capricious  in  their  choice,  rejecting  one  mare  and  without 
any  apparent  cause  taking  to  another,  that  various  artifices  have 
to  be  habitually  used.  The  famous  Monarque,  for  instance,  would 
never  consciously  look  at  the  dam  of  Gladiateur,  and  a  trick 
had  to  be  practiced.  We  can  partly  see  the  reason  why  valuable 
race-horse  stallions,  which  are  in  such  demand  as  to  be  exhausted, 
should  be  so  particular  in  their  choice.  Mr.  Blenkiron  has  never 
known  a  mare  reject  a  horse;  but  this  has  occurred  in  Mr.  Wright's 
stable,  so  that  the  mare  had  to  be  cheated.  Prosper  Lucas** 
quotes  various  statements  from  French  authorities,  and  remarks, 
"On  voit  des  etalons  qui  s'eprennent  d'une  jument,  et  negligent 
"toutes  les  autres."  He  gives,  on  the  authority  of  Baelen,  similar 
facts  in  regard  to  bulls;  and  Mr.  H.  Reeks  assures  me  that  a 
famous  short-horn  bull  belonging  to  his  father  "invariably  refused 
"to  be  matched  with  a  black  cow."  Hoffberg,  in  describing  the 
domesticated  reindeer  of  Lapland  says,  "Fcsminse  majores  et  for- 
"tiores  mares  prse  cseteris  admittunt,  ad  eos  confugiunt,  a  Juniori- 
"bus  agitatae,  qui  hos  in  fugam  conjiciunt."**  A  clergyman,  who 
has  bred  many  pigs,  asserts  that  sows  often  reject  one  boar  and 
Immediately  accept  another. 

From  these  facts  there  can  be  no  doubt  that,  with  most  of  our 
domesticated  quadrupeds,  strong  individual  antipathies  and  pref- 
erences are  frequently  exhibited,  and  much  more  commonly  by 
tHe  female  than  by  the  male.  This  being  the  case,  it  is  improb- 
able that  the  unions  of  quadrupeds  in  a  state  of  nature  should 
be  left  to  mere  chance.  It  is  much  more  probable  that  the  fe- 
males are  allured  or  excited  by  particular  males,  who  possess 
certain  characters  in  a  higher  degree  than  other  males;  but 
what  these  characters  are,  we  can  seldom  or  never  discover  with 
certainty. 

<8  'Traite  de  I'Hered.  Nat.'  torn,  ii,  1850,  p.  296, 
*»  'Amoenitates  Acad.*  vol.  iVo  1788,  p.  160. 


MAMMALS — VOCAL  ORGANS.  §21 


CHAPTER  XVIII. 

SECONDARY  SEXUAL.  CHARACTERS   OF  MAMMALS— continued. 

Voice— Remarkable  sexual  peculiarities  in  seals — Odor — Development 
of  the  hair— Color  of  the  hair  and  skin — Anomalous  case  of  the 
female  being-  more  ornamented  than  the  male— Color  and  orna- 
ments due  to  sexual  selection— Color  acquired  for  the  sake  of  pro- 
tection-Color, though  common  to  both  sexes,  often  due  to  sexual 
selection— On  the  disappearance  of  spots  and  stripes  in  adult  quad- 
rupeds—On the  colors  and  ornaments  of  the  Quadrumana— Sum- 
mary. 

Quadrupeds  use  their  voices  for  various  purposes,  as  a  signal  of 
danger,  as  a  call  from  one  member  of  a  troop  to  another,  or  from 
the  mother  to  her  lost  offspring,  or  from  the  latter  for  pr6tection 
to  their  mother;  but  such  uses  need  not  here  be  considered. 
We  are  concerned  only  with  the  difference  between  the  voices  of 
the  sexes,  for  instance  between  that  of  the  lion  and  lioness,  or  of 
the  bull  and  cow.  Almost  all  male  animals  use  their  voices  much 
more  during  the  rutting-season  than  at  any  other  time;  and 
some,  as  the  giraffe  and  porcupine,^  are  said  to  be  completely 
mute  excepting  at  this  season.  As  the  throats  (i.  e.  the  larynx 
and  thyroid  bodies-)  of  stags  periodically  become  enlarged  at  the 
beginning  of  the  breeding-season,  it  might  be  thought  that  their 
powerful  voices  must  be  somehow  of  high  importance  to  them; 
but  this  is  very  doubtful.  From  information  given  to  me  by 
two  experienced  observers,  Mr.  McNeill  and  Sir  P.  Egerton,  it 
seems  that  young  stags  under  three  years  old  do  not  roar  or 
bellow;  and  that  the  old  ones  begin  bellowing  at  the  commence- 
ment of  the  breeding-season,  at  first  only  occasionally  and  mod- 
erately, whilst  they  restlessly  wander  about  in  search  of  the  fe- 
males. Their  battles  are  prefaced  by  loud  and  prolonged  bellow- 
ing, but  during  the  actual  conflict  they  are  silent.  Animals  of  all 
kinds  which  habitually  use  their  voices  utter  various  noises  under 
any  strong  emotion,  as  when  enraged  and  preparing  to  fight; 


^  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  585. 
2  Ibid.  p.  595. 


52ii  TI-IE   DESCENT  OF  MAN. 

but  this  may  merely  be  the  result  of  nervous  excitement,  which 
leads  to  the  spasmodic  contraction  of  almost  all  the  muscles  of 
the  body,  as  when  a  man  grinds  his  teeth  and  clenches  his  tists 
in  rage  or  agony.  No  doubt  stags  challenge  each  other  to  mortal 
combat  by  bellowing;  but  those  with  the  more  powerful  voices, 
unless  at  the  same  time  the  stronger,  better-armed,  and  more 
courageous,  would  not  gain  any  advantage  over  their  rivals. 

It  is  possible  that  the  roaring  of  the  lion  may  be  of  some 
service  to  him  by  striking  terror  Into  his  adversary;  for  when 
enraged  he  likewise  erects  his  mane  and  thus  instinctively  tries 
to  make  himself  appear  as  terrible  as  possible.  But  it  can  hardly 
be  supposed  that  the  bellowing  of  the  stag,  even  if  it  be  of  ser- 
vice to  him  in  this  way,  can  have  been  important  enough  to  have 
led  to  the  periodical  enlargement  of  the  throat.  Some  writers 
suggest  that  the  bellowing  serves  as  a  call  to  the  female;  but  the 
experienced  observers  above  quoted  inform  me  that  female  deer 
do  not  search  for  the  male,  though  the  males  search  eagerly  for 
the  females  as  indeed  might  be  expected  from  what  we  know  of 
the  habits  of  other  male  quadrupeds.  The  voice  of  the  female, 
on  the  other  hand,  quickly  brings  to  her  one  or  more  stags,^  as  is 
well  known  to  the  hunters  who  in  wild  countries  imitate  her  cry. 
If  we  could  believe  that  the  male  had  the  power  to  excite  or 
allure  the  female  by  his  voice,  the  periodical  enlargement  of  his 
vocal  organs  would  be  intelligible  on  the  principle  of  sexual  selec- 
tion, together  with  inheritance  limited  to  the  same  sex  and  season; 
but  we  have  no  evidence  in  favor  of  this  view.  As  the  case 
stands,  the  loud  voice  of  a  stag  during  the  breeding-season  does 
not  seem  to  be  of  any  special  service  to  him,  either  during  his 
courtship  or  battles,  or  in  any  other  way.  But  may  we  not  be- 
lieve that  the  frequent  use  of  the  voice,  under  the  strong  excite- 
ment of  love,  jealousy,  and  rage,  continued  during  many  genera- 
tions, may  at  last  have  produced  an  inherited  effect  on,  the  vocal 
organs  of  the  stag,  as  well  as  of  other  male  animals?  This  ap- 
pears to  me,  in  our  present  state  of  knowledge,  the  most  prob- 
able view. 

The  voice  of  the  adult  male  gorilla  is  tremendous,  and  he  is 
furnished  with  a  laryngeal  sack,  as  is  the  adult  male  orang.* 
The  gibbons  rank  among  the  noisiest  of  monkeys,  and  the  Suma- 
tra species  (Hylobates  syndactylus)  is  also  furnishe(?.  with  an  air 
sack;  but  Mr.  Blyth,  who  has  had  opportunities  for  observation, 
does  not  believe  that  the  male  is  noisier  than  the  female.  Hence, 
these  latter  monkeys  probably  use  their  voices  as  a  mutual  call; 
and  this  is  certainly  the  case  with  some  quadrupeds,  for  instance 


3  See,  for  instance,  Major  W.  Ross  King-  ('The  Sportsman  in  Canada,' 
1866,  p.  53,  131)  on  the  habits  of  the  moose  and  wild  reindeer. 
-  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  600. 


MAMMALS-VOCAL  ORGANS.  523 

the  beaver.^  Another  gibbon,  the  H.  agilis,  is  remarkable,  from 
having  the  power  of  giving  a  complete  and  correct  octave  of 
musical  notes,^  which  we  may  reasonably  suspect  serves  as  a 
sexual  charm;  but  I  shall  have  to  recur  to  this  subject  in  the  next 
chapter.  The  vocal  organs  of  the  American  Mycetes  caraya  are 
one-third  larger  in  the  male  than  in  the  female,  and  are  won- 
derfully powerful.  These  monkeys  in  warm  weather  make  the 
forests  resound  at  morning  and  evening  with  their  overwhelming 
voices.  The  males  begin  the  dreadful  concert,  and  often  continue 
it  during  many  hours,  the  females,  sometimes  Joining  in  with 
their  less  powerful  voices.  An  excellent  observer,  Rengger,^  could 
not  perceive  that  they  were  excited  to  begin  by  any  special  cause; 
he  thinks  that,  like  many  birds,  they  delight  in  their  own  music, 
and  try  to  excel  each  other.  Whether  most  of  the  foregoing 
monkeys  have  acquired  their  powerful  voices  in  order  to  beat 
their  rivals  and  charm  the  females — or  whether  the  vocal  organs 
have  been  strengthened  and  enlarged  through  the  inherited  ef- 
fects of  long-continued  use  without  any  particular  good  being  thus 
gained — I  will  not  pretend  to  say;  but  the  former  view,  at  least 
in  the  case  of  the  Hylobates  agilis,  seems  the  most  probable. 

I  may  here  mention  two  very  curious  sexual  peculiarities  oc- 
curring in  seals,  because  they  have  been  supposed  by  some  writ- 
ers to  affect  the  voice.  The  nose  of  the  male  sea-elephant  (Ma- 
crorhinus  proboscideus)  becomes  greatly  elongated  during  the 
breeding-season,  and  can  then  be  erected.  In  this  state  it  is 
sometimes  a  foot  in  length.  The  female  is  not  thus  provided  at 
any  period  of  life.  The  male  makes  a  wild,  hoarse,  gurgling 
noise,  which  is  audible  at  a  great  distance  and  is  believed  to  be 
strengthened  by  the  proboscis;  the  voice  of  the  female  being 
different.  Lesson  compares  the  erection  of  the  proboscis,  with 
the  swelling  of  the  wattles  of  male  gallinaceous  birds  whilst 
courting  the  females.  In  another  allied  kind  of  seal,  the  bladder- 
nose  (Cystophora  cristata),  the  head  is  covered  by  a  great  hood 
or  bladder.  This  is  supported  by  the  septum  of  the  nose,  which 
is  produced  far  backwards  and  rises  into  an  internal  crest  seven 
inches  in  height.  The  hood  is  clothed  with  short  hair,  and  is 
muscular;  it  can  be  inflated  until  it  more  than  equals  the  whole 
head  in  size!  The  males  when  rutting,  fight  furiously  on  the 
ice,  and  their  roaring  "is  said  to  be  sometimes  so  loud  as  to 
"be  heard  four  miles  off."  When  attacked  they  likewise  roar  or 
bellow;  and  whenever  irritated  the  bladder  is  inflated  and  quiv- 
ers.    Some  naturalists  believe  that  the  voice  is  thus  strengthened. 


^  Mr.  Green,  in  'Journal  of  Linn.  Soc'  vol.  x.  Zoology,  1869,  p.  362. 
6  C.   L.  Martin,    'General  Introduction  to  the  Nat.   Hist,   of  Mamm. 
Animals,'  1841,  p.  431. 
'  Naturgeschichte  der  Saugethiere  von  Paraguay,'  1830,  s.  15,  21. 


524  THE  DESCENT  OF  MAN. 

but  various  other  uses  have  been  assigned  to  this  extraordinary 
structure.  Mr.  R.  Brown  thinks  that  it  serves  as  a  protection 
against  accidents  of  all  kinds;  but  this  is  not  probable,  for,  as  I 
am  assured  by  Mr.  Lamont  who  killed  600  of  these  animals,  the 
hood  is  rudimentary  in  the  females,  and  it  is  not  developed  in  the 
males  during  youth.^ 

Qcior^ — With  some  animals,  as  with  the  notorious  skunk  of 
America,  the  overwhelming  odor  which  they  emit  appears  to 
serve  exclusively  as  a  defense.  With  shrew-mice  (Sorex)  both 
sexes  possess  abdominal  scent-glands,  and  there  can  be  little 
doubt,  from  the  rejection  of  their  bodies  by  birds  and  beasts  of 
prey,  that  the  odor  is  protective;  nevertheless,  the  glands  be- 
come enlarged  in  the  males  during  the  breeding-season.  In  many 
other  quadrupeds  the  glands  are  of  the  same  size  in  both  sexes,^ 
but  their  uses  are  not  known.  In  other  species  the  glands  are 
confined  to  the  males,  or  are  more  developed  than  in  the  females; 
and  they  almost  always  become  more  active  during  the  rutting- 
season.  At  this  period  the  glands  on  the  sides  of  the  face  of  the 
male  elephant  enlarge,  and  emit  a  secretion  having  a  strong 
musky  odor.  The  males,  and  rarely  the  females,  of  many  kinds 
of  bats  have  glands  and  protrudable  sacks  situated  in  various 
parts;    and  it  is  believed  that  these  are  odoriferous. 

The  rank  effluvium  of  the  male  goat  is  well  known,  and  that 
of  certain  male  deer  is  wonderfully  strong  and  persistent.  On 
the  banks  of  the  Plata  I  perceived  the  air  tainted  with  the  odor 
of  the  male  Cervus  campestris,  at  half  a  mile  to  leeward  of  a 
herd;  and  a  silk  handkerchief,  in  which  I  carried  home  a  skin, 
though  often  used  and  washed,  retained,  when  first  unfolded, 
traces  of  the  odor  for  one  year  and  seven  months.  This  animal 
does  not  emit  its  strong  odor  until  more  than  a  year  old,  and  if 
castrated  whilst  young  never  emits  it.^°    Besides  the  general  odor. 


8  On  the  sea-elephant,  see  an  article  by  Lesson,  in  'Diet.  Class.  Hist. 
Nat.'  torn.  xiii.  p.  418.  For  the  Cystophora  or  Stemmatopus,  see  Dr. 
Dekay,  'Annals  of  Lyceum  of  Nat.  Hist.  New  York,'  vol.  i.  1824,  p.  94. 
Pennant  has  also  collected  information  from  the  sealers  on  this  ani- 
mal. The  fullest  account  is  given  by  Mr.  Brown,  in  'Proc.  Zoolog. 
Soc'  1868,  p.  435. 

»  As  with  the  castoreum  of  the  beaver,  see  Mr.  L.  H.  Morgan's  most 
interesting  work,  'The  American  Beaver,'  1868,  p.  300.  Pallas  ('Spic. 
Zoolog.'  fasc.  viii.  1779,  p.  23)  has  well  discussed  the  odoriferous  glands 
of  mammals.  Owen  ('Anat.  of  Vertebrates,'  vol.  iii.  p.  634)  also  gives 
an  account  of  these  glands,  including  those  of  the  elephant,  and 
(p.  763)  those  of  shrew-mice.  On  Bats,  Mr.  Dobson,  in  'Proc.  Zoolog. 
Soc'  1873,  p.  241. 

^°  Rengger,  'Naturgeschichte  der  Saugethiere  von  Paraguay,'  1830, 
s.  355.  This  observer  also  gives  some  curious  particulars  in  regard  to 
the  odor. 


MAMMALS— ODORS  EMITTED.  525 

permeating  the  whole  body  of  certain  ruminants  (for  instance. 
Bos  moschatus)  in  the  breeding-season,  many  deer,  antelopes, 
sheep,  and  goats,  possess  odoriferous  glands  in  various  situations, 
more  especially  on  their  faces.  The  so-called  tear-sacks,  or  sub- 
orbital pits,  come  under  this  head.  These  glands  secrete  a  semi- 
fluid fetid  matter  which  is  sometimes  so  copious  as  to  stain  the 
whole  face,  as  I  have  myself  seen  in  an  antelope.  They  are 
"usually  larger  in  the  male  than  in  the  female,  and  their  devel- 
"opment  is  checked  by  castration."^^  According  to  Desmarest  they 
are  altogether  absent  in  the  female  of  Antilope  subgutturosa. 
Hence,  there  can  be  no  doubt  that  they  stand  in  close  relation  with 
the  reproductive  functions.  They  are  also  sometimes  present, 
and  sometimes  absent,  in  nearly-allied  forms.  In  the  adult  male 
musk-deer  (Moschus  moschiferus),  a  naked  space  round  the  tail 
is  bedewed  with  an  odoriferous  fluid,  whilst  in  the  adult  female, 
and  in  the  male  until  two  years  old,  this  space  is  covered  with 
hair  and  is  not  odoriferous.  The  proper  musk-sack  of  this  deer 
is  from  its  position  necessarily  conflned  to  the  male,  and  forms 
an  additional  scent-organ.  It  is  a  singular  fact  that  the  matter 
secreted  by  the  latter  gland  does  not,  according  to  Pallas,  change 
in  consistence,  or  increase  in  quantity,  during  the  rutting-season; 
nevertheless  this  naturalist  admits  that  its  presence  is  in  some 
way  connected  with  the  act  of  reproduction.  He  gives,  however, 
only  a  conjectural  and  unsatisfactory  explanation  of  its  use.^ 

In  most  cases,  when  only  the  male  emits  a  strong  odor  during 
the  breeding-season,  it  probably  serves  to  excite  or  allure  the  fe- 
male. We  must  not  judge  on  this  head  by  our  own  taste,  for  it 
is  well  known  that  rats  are  enticed  by  certain  essential  oils,  and 
cats  by  valerian,  substances  far  from  agreeable  to  us;  and  that 
dogs,  though  they  will  not  eat  carrion,  sniff  and  roll  on  it.  From 
the  reasons  given  when  discussing  the  voice  of  the  stag,  we  may 
reject  the  idea  that  the  odor  serves  to  bring  the  females  from  a 
distance  to  the  males.  Active  and  long-continued  use  cannot 
here  have  come  into  play,  as  in  the  case  of  the  vocal  organs.  The 
odor  emitted  must  be  of  considerable  importance  to  the  male, 
inasmuch  as  large  and  complex  glands,  furnished  with  muscles 
for  everting  the  sack,  and  for  closing  or  opening  the  orifice,  have 
in  some  cases  been  developed.  The  development  of  these  organs 
is  intelligible  through  sexual  selection,  if  the  most  odoriferous 
males  are  the  most  successful  in  winning  the  females,  and  in 

^  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii.  p.  632.  See,  also,  Dr. 
Murie's  observations  on  these  glands  in  the  'Proc.  Zoolog.  Soc'  1870, 
p.  340.  Desmarest,  on  the  Antilope  subgutturosa,  'Mammalogie,'  1820, 
p.  455. 

^  Pallas,  'Spicilegia  Zoolog.'  fasc.  xiii.  1799,  p.  24;  Desmoulins,  'Diet. 
Class,  d'Hist.  Nat.'  torn.  iii.  p.  586. 


526  THE  DESCENT  OF  MAN. 

leaving  offspring  to  inherit  their  gradually-perfected  glands  and 
odors. 

Development  of  the  Hair.  — We  have  seen  that  male  quadrupeds 
often  have  the  hair  on  their  necks  and  shoulders  much  more 
developed  than  the  females;  and  many  additional  instances  could 
be  given.  This  sometimes  serves  as  a  defense  to  the  male  during 
his  battles;  but  whether  the  hair  in  most  cases  has  been  specially 
developed  for  this  purpose,  is  very  doubtful.  We  may  feel  al- 
most certain  that  this  is  not  the  case,  when  only  a  thin  and 
narrow  crest  runs  along  the  back;  for  a  crest  of  this  kind  would 
afford  scarcely  any  protection,  and  the  ridge  of  the  back  is  not 
a  place  likely  to  be  injured;  nevertheless  such  crests  are  some- 
times confined  to  the  males,  or  are  much  more  developed  in  them 
than  in  the  females.  Two  antelopes,  the  Tragelaphus  scriptus"^' 
(see  fig.  70,  p.  539)  and  Portax  picta,  may  be  given  as  instances. 
When  stags,  and  the  males  of  the  wild  goat,  are  enraged  or  terri- 
fied, these  crests  stand  erect;'*  but  it  cannot  be  supposed  that  they 
have  been  developed  merely  for  the  sake  of  exciting  fear  in  their 
enemies.  One  of  the  above-named  antelopes,  the  Portax  picta, 
has  a  large  well-defined  brush  of  black  hair  on  the  throat,  and 
this  is  much  larger  in  the  male  than  in  the  female.  In  the  Am- 
motragus  tragelaphus  of  North  Africa,  a  member  of  the  sheep- 
family,  the  fore-legs  are  almost  concealed  by  an  extraordinary 
growth  of  hair,  which  depends  from  the  neck  and  upper  halves  of 
the  legs;  but  Mr.  Bartlett  does  not  believe  that  this  mantle  is 
of  the  least  use  to  the  male,  in  whom  it  is  much  more  developed 
than  in  the  female. 

Male  quadrupeds  of  many  kinds  differ  from  the  females  in  hav- 
ing more  hair,  or  hair  of  a  different  character,  on  certain  parts 
of  their  faces.  Thus  the  bull  alone  has  curled  hair  on  the  fore- 
head.^^  In  three  closely-allied  sub-genera  of  the  goat  family, 
only  the  males  possess  beards,  sometimes  of  large  size;  in  two 
other  sub-genera  both  sexes  have  a  beard,  but  it  disappears  in 
some  of  the  domestic  breeds  of  the  common  goat;  and  neither 
sex  of  the  Hemitragus  has  a  beard.  In  the  ibex  the  beard  is  not 
developed  during  the  summer,  and  it  is  so  small  at  other  times 
that  it  may  be  called  rudimentary.'^  With  some  monkeys  the 
beard  is  confined  to  the  male,  as  in  the  orang;    or  is  much  larger 


13  Dr.  Gray,  'Gleanings  from  the  Menagerie  at  Knowsley,'  pi.  28. 

1*  Judge  Caton  on  the  Wapiti,  'Transact.  Ottawa  Acad.  Nat.  Sci- 
ences,' 1868,  pp.  36,  40;  Blyth,  'Land  and  Water,'  on  Capra  aegagrus, 
1867,  p.  37. 

15  'Hunter's  Essays  and  Observations,'  edited  by  Owen,  1861,  vol.  i. 
p.  236. 

18  See  Dr.  Gray's  'Cat.  of  Mammalia  in  British  Museum/  part  iii. 
1852,  p.  144. 


MAMMALS— DEVELOPMENT  OF  HAIR. 


527 


in  the  male  than  in  the  female,  as  in  the  Mycetes  caraya  and 
Pithecia  satanas  (fig.  68).  So  it  is  with  the  whiskers  of  some 
species  of  Macacus,"  and,  as  we  have  seen,  with  the  manes  of 
some  species  of  baboons.  But  with  most  kinds  of  monkeys  the 
various  tufts  of  hair  about  the  face  and  head  are  alike  in  both 
sexes. 

The  males  of  various  members  of  the  ox  family  (Bovidse),  and 
of  certain  antelopes,  are  furnished  with  a  dewlap,  or  great  fold  of 
skin  on  the  neck,  which  is  much  less  developed  in  the  female. 


Fig-.  68.    Pithecia  satanas,  male  (from  Brehm). 

Now,  what  must  we  conclude  with  respect  to  such  sexual  dif- 
ferences as  these?  No  one  will  pretend  that  the  beards  of  cer- 
tain male  goats,  or  the  dewlap  of  the  bull,  or  the  crests  of  hair 
along  the  backs  of  certain  male  antelopes,  are  of  any  use  to  them 
in  their  ordinary  habits.  It  is  possible  that  the  immense  beard 
of  the  male  Pithecia,  and  the  large  beard  of  the  male  orang, 
may  protect  their  throats  when  fighting;  for  the  keepers  in  the 
Zoological  Gardens  inform  me  that  many  monkeys  attack  each 


"  Rengger,  'Saugethiere,'  «&;c.  s.  14;    Desmarest,  'Mammalogie,'  p.  66. 


528  THE  DESCENT  OF  MAN. 

other  by  the  throat;  but  it  is  not  probable  that  the  beard  has 
been  developed  for  a  distinct  purpose  from  that  served  by  the 
whiskers,  moustache,  and  other  tufts  of  hair  on  the  face;  and 
no  one  will  suppose  that  these  are  useful  as  a  protection.  Must 
we  attribute  all  these  appendages  of  hair  or  skin  to  mere  pur- 
poseless variability  in  the  male?  It  cannot  be  denied  that  this 
is  possible;  for  in  many  domesticated  quadrupeds,  certain  char- 
acters, apparently  not  derived  through  reversion  from  any  wild 
parent-form,  are  confined  to  the  males,  or  are  more  developed  in 
them  than  in  the  females — for  instance,  the  hump  on  the  male 
zebu-cattle  of  India,  the  tail  of  fat-tailed  rams,  the  arched  out- 
line of  the  forehead  in  the  males  of  several  breeds  of  sheep,  and, 
lastly,  the  mane,  the  long  hairs  on  the  hind-legs,  and  the  dewlap 
of  the  male  of  the  Berbura  goat.^^  The  mane,  which  occurs  only 
in  the  rams  of  an  African  breed  of  sheep,  is  a  true  secondary  sex- 
ual character,  for,  as  I  hear  from  Mr.  Winwood  Reade,  it  is  not 
developed  if  the  animal  be  castrated.  Although  we  ought  to  be 
extremely  cautious,  as  shown  in  my  work  on  'Variation  under 
Domestication,'  in  concluding  that  any  character,  even  with  ani- 
mals kept  by  semi-civilized  people,  has  not  been  subjected  to 
selection  by  man,  and  thus  augmented,  yet  in  the  cases  just  speci- 
fied this  is  improbable;  more  especially  as  the  characters  are 
confined  to  the  males,  or  are  more  strongly  developed  in  them 
than  in  the  females.  If  it  were  positively  known  that  the  above 
African  ram  is  a  descendant  of  the  same  primitive  stock  as  the 
other  breeds  of  sheep,  and  if  the  Berbura  male-goat  with  his 
mane,  dewlap,  &c.,  is  descended  from  the  same  stock  as  other 
goats,  then,  assuming  that  selection  has  not  been  applied  to  these 
characters,  they  must  be  due  to  simple  variability,  together  with 
sexually-limited  inheritance. 

Hence  it  appears  reasonable  to  extend  this  same  view  to  all 
analogous  cases  with  animals  in  a  state  of  nature.  Nevertheless 
I  cannot  persuade  myself  that  it  generally  holds  good,  as  in  the 
case  of  the  extraordinary  development  of  hair  on  the  throat  and 
fore-legs  of  the  male  Ammotragus,  or  in  that  of  the  immense 
beard  of  the  male  Pithecia.  Such  study  as  I  have  been  able  to 
give  to  nature  makes  me  believe  that  parts  or  organs  which  are 
highly  developed,  were  acquired  at  some  period  for  a  special 
purpose.  With  those  antelopes  in  which  the  adult  male  is  more 
strongly-colored  than  the  female,  and  with  those  monkeys  in 
which  the  hair  on  the  face  is  elegantly  arranged  and  colored  in 
a  diversified  manner,  it  seems  probable  that  the  crests  and  tufts 

18  See  the  chapters  on  these  several  animals  in  vol.  i.  of  my  'Varia- 
tion of  Animals  under  Domestication;'  also,  vol.  ii.  p.  73;  also,  chap. 
XX.  on  the  practice  of  selection  by  semi-civilized  people.  For  the  Ber- 
bura goat,   see  Dr.  Gray,  'Catalogue,'  ibid.  p.  157. 


MAMMALS— ORNAMENTAL   COLORS.  529 

of  hair  were  gained  as  ornaments;  and  this  I  know  Is  the  opinion 
of  some  naturalists.  If  this  be  correct,  there  can  be  little  doubt 
that  they  were  gained  or  at  least  modified  through  sexual  selec- 
tion; but  how  far  the  same  view  may  be  extended  to  other  mam- 
mals is  doubtful. 

Coliyr  of  the  Hair  and  of  the  Naked  Skin. — I  will  first  give 
briefiy  all  the  cases  known  to  me,  of  male  quadrupeds  differing 
in  color  from  the  females.  "With  Marsupials,  as  I  am  informed 
by  Mr.  Gould,  the  sexes  rarely  differ  in  this  respect;  but  the 
great  red  kangaroo  offers  a  striking  exception,  "delicate  blue 
"being  the  prevailing  tint  in  those  parts  of  the  female,  which 
"in  the  male  are  red."^®  In  the  Didelphis  opossum  of  Cayenne 
the  female  is  said  to  be  a  little  more  red  than  the  male.  Of  the 
Rodents  Dr.  Gray  remarks:  "African  squirrels,  especially  those 
"found  in  the  tropical  regions,  have  the  fur  much  brighter  and 
"more  vivid  at  some  seasons  of  the  year  than  at  others,  and  the 
"fur  of  the  male  is  generally  brighter  than  that  of  the  female.''^^" 
Dr.  Gray  informs  me  that  "he  specified  the  African  squirrels,  be- 
cause, from  their  unusually  bright  colors,  they  best  exhibit  this 
difference.  The  female  of  the  Mus  minutus  of  Russia  is  of  a  paler 
and  dirtier  tint  than  the  male.  In  a  large  number  of  bats  the 
fur  of  the  male  is  lighter  than  in  the  female.^^  Mr.  Dobson  also 
remarks,  with  respect  to  these  animals:  "Differences,  depending 
"partly  or  entirely  on  the  possession  by  the  male  of  fur  of  a  much 
"more  brilliant  hue,  or  distinguished  by  different  markings  or 
"by  the  greater  length  of  certain  portions,  are  met  only,  to  any 
"appreciable  extent,  in  the  frugivorous  bats  in  which  the  sense 
"of  sight  is  well  developed."  This  last  remark  deserves  attention, 
as  bearing  on  the  question  whether  bright  colors  are  serviceable 
to  male  animals  from  being  ornamental.  In  one  genus  of 
sloths,  it  is  now  established,  as  Dr.  Gray  states,  "that  the  males 
"are  ornamented  differently  from  the  females — that  is  to  say, 
"that  they  have  a  patch  of  soft  short  hair  between  the  shoulders, 
"which  is  generally  of  a  more  or  less  orange  color,  and  in  one 
"species  pure  white.  The  females  on  the  contrary,  are  destitute  of 
"this  mark." 

The  terrestrial  Carnivora  and  Insectivora  rarely  exhibit  sexual 
differences  of  any  kind,  including  color.    The  ocelot  (Felis  par- 

i»  Osphranter  rufus,  Gould,  'Mammals  of  Australia,'  1863,  vol.  ii.    On 
the  Didelphis,  Desmarest,  'Mammalogie,'  p.  256. 

2"  'Annals  and  Mag.   of  Nat.   Hist.'   Nov.  1867,   p.  325.    On    the   Mus 
minutus,  Desmarest,  'Mammalogie,'  p.  304. 

21  J.   A.   Allen,    in    'Bulletin   of  Mus.    Comp.    Zoolog-.    of   Cambridge, 
United  States,'  1869,  p.  207.    Mr.  Dobson  on  sexual  characters  in  the 
Chiroptera,  'Proc.  Zoolog.  Soc'  1873,  p.  241.    Dr.  Gray  on  Sloths,  ibid. 
1871,  p.  436. 
35 


530  THE  DESCENT  OP  MAN. 

dalis),  however,  is  exceptional,  for  the  colors  of  the  female,  com- 
pared with  those  of  the  male,  are  "moins  apparentes,  le  fauve 
"etant  plus  terne,  le  blanc  moins  pur,  les  raies  ayant  moins  de 
"largeur  et  les  taches  moins  de  diametre."-  The  sexes  of  the 
allied  Felis  mitis  also  differ,  but  in  a  less  degree;  the  general 
hues  of  the  female  being  rather  paler  than  in  the  male,  with 
the  spots  less  black.  The  marine  Carnivora  or  seals,  on  the  other 
hand,  sometimes  differ  considerably  in  color,  and  they  present, 
as  we  have  already  seen,  other  re'markable  sexual  differences. 
Thus  the  male  of  the  Otaria  nigrescens  of  the  southern  hemi- 
sphere is  of  a  rich  brown  shade  above;  whilst  the  female,  who  ac- 
quires her  adult  tints  earlier  in  life  than  the  male,  is  dark-gray 
above,  the  young  of  both  sexes  being  of  a  deep  chocolate  color.  The 
male  of  the  northern  Phoca  groenlandica  is  tawny  gray,  with  a 
curious  saddle-shaped  dark  mark  on  the  back;  the  female  is 
much  smaller,  and  has  a  very  different  appearance,  being  "dull 
"white  or  yellowish  straw-color,  with  a  tawny  hue  on  the  back;" 
the  young  at  first  are  pure  white,  and  can  "hardly  be  distinguished 
"among  the  icy  hummocks  and  snow,  their  color  thus  acting  as  a 
"protection."-^ 

With  Ruminants  sexual  differences  of  color  occur  more  com- 
monly than  in  any  other  order.  A  difference  of  this  kind  is  gen- 
eral in  the  Strepsicerene  antelopes;  thus  the  male  nilghau  (Por- 
tax  picta)  is  bluish-gray  and  much  darker  than  the  female,  with 
the  square  white  patch  on  the  throat,  the  white  marks  on  the 
fetlocks,  and  the  black  spots  on  the  ears  all  much  more  distinct. 
We  have  seen  that  in  this  species  the  crests  and  tufts  of  hair  are 
likewise  more  developed  in  the  male  than  in  the  hornless  female. 
I  am  informed  by  Mr.  Blyth  that  the  male,  without  shedding 
his  hair,  periodically  becomes  darker  during  the  breeding-season. 
Young  males  cannot  be  distinguished  from  young  females  until 
about  twelve  months  old;  and  if  the  male  is  emasculated  before 
this  period,  he  never,  according  to  the  same  authority,  changes 
color.  The  importance  of  this  latter  fact,  as  evidence  that  the 
coloring  of  the  Portax  is  of  sexual  origin  becomes  obvious,  when 
we  hear^*  that  neither  the  red  summer-coat  nor  the  blue  winter- 
coat  of  the  Virginian  deer  is  at  all  affected  by  emasculation.  With 
most  or  all  of  the  highly-ornamented  species  of  Tragelaphus  the 
males  are  darker  than  the  hornless  females,  and  their  crests  of 


22Desmarest,  'Mammalogie,'  1820,  p.  220.  On  Felis  mitis  Rengger, 
ibid.  s.  194. 

23  Dr.  Murie  on  the  Otaria,  'Proc.  Zool.  Soc'  1869,  p.  108.  Mr.  R. 
Brown,  on  the  P.  groenlandica,  ibid.  1868,  p.  417.  See,  also,  on  the  colors 
Of  seals,  Desmarest,  ibid.  p.  243,  249. 

*'  Judge  Caton,  in  'Trans.  Ottawa  Acad,  of  Nat.  Sciences,'  1868,  p.  4. 


MAMMALS— ORNAMENTAL  COLORS.  531 

hair  are  more  fully  developed.  In  the  male  of  that  magnificent 
antelope,  the  Derbyan  eland,  the  body  is  redder,  the  whole  neck 
much  blacker,  and  the  white  band  which  separates  these  colors, 
broader,  than  in  the  female.  In  the  Cape  eland  also,  the  male  is 
slightly  darker  than  the  female.^^ 

In  the  Indian  black-buck  (A.  bezoartica),  which  belongs  to  an- 
other tribe  of  antelopes,  the  male  is  very  dark,  almost  black; 
whilst  the  hornless  female  is  fawn-colored.  We  meet  in  this 
species,  as  Mr.  Blyth  informs  me,  with  an  exactly  similar  series 
of  facts,  as  in  the  Portax  picta,  camely,  in  the  male  periodically 
changing  color  during  the  breeding-season,  in  the  effects  of  emas- 
culation on  this  change,  and  in  the  young  of  both  sexes  being 
indistinguishable  from  each  other.  In  the  Antilope  niger  the 
male  is  black,  the  female,  as  well  as  the  young  of  both  sexes, 
being  brown;  in  A.  sing-sing  the  male  is  much  brighter  colored 
than  the  hornless  female,  and  his  chest  and  belly  are  blacker; 
in  the  male  A.  caama,  the  marks  and  lines  which  occur  on  various 
parts  of  the  body  are  black,  instead  of  brown  as  in  the  female; 
in  the  brindled  gnu  (A.  gorgon)  "the  colors  of  the  male  are 
"nearly  the  same  as  those  of  the  female,  only  deeper  and  of  a 
"brighter  hue."^°    Other  analogous  cases  could  be  added. 

The  Banteng  bull  (Bos  sondaicus)  of  the  Malayan  Archipelago 
is  almost  black,  with  white  legs  and  buttocks;  the  cow  is  of  a 
bright  dun,  as  are  the  young  males  until  about  the  age  of  three 
years,  when  they  rapidly  change  color.  The  emasculated  bull 
reverts  to  the  color  of  the  female.  The  female  Kemas  goat  is 
paler,  and  both  it  and  the  female  Capra  segagrus  are  said  to  be 
more  uniformly  tinted  than  their  males.  Deer  rarely  present 
any  sexual  differences  in  color.  Judge  Caton,  however,  informs 
me  that  in  the  males  of  the  wapiti  deer  (Cervus  canadensis)  the 
neck,  belly,  and  legs  are  much  darker  than  in  tlie  female;  but 
during  the  winter  the  darker  tints  gradually  fade  away  and  dis- 
appear. I  may  here  mention  that  Judge  Caton  has  in  his  park 
three  races  of  the  Virginian  deer,  which  differ  slightly  in  color, 
but  the  differences  are  almost  exclusively  confined  to  the  blue 

2^  Dr.  Gray,  'Cat.  of  Manim.  in  Brit.  Mus.'  part  iil.  1852,  pp.  134-142; 
also,  Dr.  Gray,  'Gleanings  from  the  Menagerie  of  Knowsley,'  in  which 
there  is  a  splendid  drawing  of  the  Oreas  derhianus:  see  the  text  on 
Tragelaphus.  For  the  Cape  eland  (Oreas  canna),  see  Andrew  Smith, 
'Zoology  of  S.  Africa,'  pi.  41  and  42.  There  are  also  many  of  these 
antelopes  in  the  Zoological  Gardens. 

26  On  the  Ant.  niger,  see  'Proc.  Zool.  Soc'  1850,  p.  133.  With  respect 
to  an  allied  species,  in  which  there  is  an  equal  sexual  difference  in 
color,  see  Sir  S.  Baker,  'The  Albert  Nyanza,'  1866,  vol.  ii.  p.  327.  For 
the  A.  sing-sing,  Gray,  'Cat.  B.  Mus.'  p.  100.  Desmarest,  'Mammalo- 
gie,'  p.  468,  on  the  A.  caama.  Andrew  Smith,  'Zoology  of  S.  Africa,' 
on  the  Gnu. 


532  THE  DESCENT  OF  MAN. 

winter  or  breeding  coat;  so  that  tbis  case  may  be  compared  with 
those  given  in  a  previous  chapter  of  closely-allied  or  representa- 
tive species  of  birds,  which  differ  from  each  other  only  in  their 
breeding  plumage.-^  The  females  of  Cervus  paludosus  of  S.  Amer- 
ica, as  well  as  the  young  of  both  sexes,  do  not  possess  the  black 
stripes  on  the  nose  and  the  blackish-brown  line  on  the  breast, 
which  are  characteristic  of  the  adult  males.-*  Lastly,  as  I  am  in- 
formed by  Mr.  Blyth,  the  mature  male  of  the  beautifully  colored 
and  spotted  axis  deer  is  considerably  darker  than  the  female;  and 
this  hue  the  castrated  male  never  acquires. 

The  last  Order  which  we  need  consider  is  that  of  the  Primates. 
The  male  of  the  Lemur  macaco  is  generally  coal-black,  whilst  the 
female  is  brown.-"  Of  the  Quadrumana  of  the  New  World,  the 
females  and  young  of  Mycetes  caraya  are  grayish-yellow  and  like 
each  other;  in  the  second  year  the  young  male  becomes  reddish- 
brown;  in  the  third,  black,  excepting  the  stomach,  which,  how- 
ever, becomes  quite  black  in  the  fourth  or  fifth  year.  There  is 
also  a  strongly-marked  difference  in  color  between  the  sexes  ol 
Mycetes  seniculus  and  Cebus  capucinus;  the  young  of  the  former, 
and  I  believe  of  the  latter  species,  resembling  the  females.  With 
Pithecia  leucocephala  the  young  likewise  resemble  the  females, 
which  are  brownish-black  above  and  light  rusty-red  beneath,  the 
adult  males  being  black.  The  rulf  of  hair  round  the  face  of  Ateles 
marginatus  is  tinted  yellow  in  the  male  and  white  in  the  female. 
Turning  to  the  Old  World,  the  males  of  Hylobates  hoolock  are 
always  black,  with  the  exception  of  a  white  band  over  the  brows; 
the  females  vary  from  whity-brown  to  a  dark  tint  mixed  with 
black,  but  are  never  wholly  black.^°  In  the  beautiful  Cercopithe- 
cus  diana,  the  head  of  the  adult  male  is  of  an  intense  black,  whilst 
that  of  the  female  is  dark  gray;  in  the  former  the  fur  between 
the  thighs  is  of  an  elegant  fawn-color,  in  the  latter  it  is  paler. 
In  the  beautiful  and  curious  moustache  monkey  (Cercopithecus 
cephus)  the  only  difference  between  the  sexes  is  that  the  tail  of 
the  male  is  chestnut  and  that  of  the  female  gray;    but  Mr.  Bart- 

27  'Ottawa  Academy  of  Sciences,'  May  21,  1868,  pp.  3,  5. 

28  S.  Muller,  on  the  Banteng,  'Zoog-.  Indischen  Archipel.'  1839-1844. 
tab.  35;  see,  also,  Raffles,  as  quoted  by  Mr.  Blyth,  in  'Land  and 
Water,'  1867,  p.  476.  On  goats,  Dr.  Gray,  'Cat.  Brit.  Mus.'  p.  146;  Des- 
marest,  'Mammalogie,'  p.  482.  On  the  Cervus  paludosus,  Rengger,  ibid, 
s.  345. 

2»  Sclater,  'Proc.  Zool.  Soc'  1866,  p.  1.  The  same  fact  has  also  been 
fully  ascertained  by  MM.  Pollen  and  van  Dam.  See,  also,  Dr.  Gray 
in  'Annals  and  Mag.  of  Nat.  Hist.'  May,  1871,  p.  340. 

30  On  Mycetes,  Rengger,  ibid.  s.  14;  and  Brehm,  'Illustrirtes  Thier- 
leben,'  B.  1.  s.  96,  107.  On  Ateles,  Desmarest,  'Mammalogie,'  p.  75.  On 
Hylobates,  Blyth,  'Land  and  Water,'  1867,  p.  135.  On  the  Semnoplthe- 
cus,  S.  Muller,  'Zoog.  Indischen  Archipel,'  tab.  x. 


MAMMALS— ORNAMENTAL  COLORS. 


533 


lett  informs  me  that  all  the  hues  become  more  pronounced  in  the 
male  when  adult,  whilst  in  the  female  they  remain  as  tney  were 
during  youth.  According  to  the  colored  figures  given  by  Solomon 
Miiller,  the  male  of  Semnopithecus  chrysomelas  is  nearly  black, 
the  female  being  pale  brown.  In  the  Cercopithecus  cynosurus  and 
griseoviridis  one  part  of  the  body,  which  is  confined  to  the  male 
sex,  is  of  the  most  brilliant  blue  or  green,  and  contrasts  strikingly 


Fig.  69.    Head  of  Male  Mandrill  (from  Gervais,  'Hist.  Nat.  des  Mam- 

miferes'). 

with  the  naked  skin  on  the  hinder  part  of  the  body,  which  is 
vivid  red. 

Lastly,  in  the  baboon  family,  the  adult  male  of  Cynocephalus 
hamadryas  differs  from  the  female  not  only  by  his  immense  mane, 
but  slightly  in  the  color  of  the  hair  and  of  the  naked  callosities. 
In  the  drill  (C.  leucophseus)  the  females  and  young  are  much 
paler-colored,  with  less  green,  than  the  adult  males.  No  other 
member  in  the  whole  class  of  mammals  is  colored  in  so  extraor- 
36 


534  THE  DESCENT  OP  MAN. 

dinary  a  manner  as  the  adult  male  mandrill  (C.  mormon).  The 
face  at  this  age  becomes  of  a  tine  blue,  with  the  ridge  and  tip 
of  the  nose  of  the  most  brilliant  red.  According  to  some  authors, 
the  face  is  also  marked  with  whitish  stripes,  and  is  shaded  in 
parts  with  black,  but  the  colors  appear  to  be  variable.  On  the 
forehead  there  is  a  crest  of  hair,  and  on  the  chin  a  yellow  beard. 
"Toutes  les  parties  superieures  de  leurs  cuisses  et  le  grand  espace 
*'nu  de  leurs  fesses  sont  ogalement  colores  du  rouge  le  plus  vif, 
^'avec  un  melahge  de  bleu  qui  ne  manque  reellement  pas 
"d'elegance."^^  When  the  animal  is  excited  all  the  naked  parts 
become  much  more  vividly  tinted.  Several  authors  have  used  the 
strongest  expressions  in  describing  these  resplendent  colors, 
which  they  compare  with  those  of  the  most  brilliant  birds.  An- 
other remarkable  peculiarity  is  that  when  the  great  canine  teeth 
are  fully  developed,  immense  protuberances  of  bone  are  formed 
on  each  cheek,  which  are  deeply  furrowed  longitudinally,  and 
the  naked  skin  ovfer  them  is  brilliantly-colored,  as  just  described. 
(Fig.  69.)  In  the  adult  females  and  in  the  young  of  both  sexes 
these  protuberances  are  scarcely  perceptible;  and  the  naked  parts 
are  much  less  bright  colored,  the  face  being  almost  black,  tinged 
with  blue.  In  the  adult  female,  however,  the  nose  at  certain  regu- 
lar intervals  of  time  becomes  tinted  with  red. 

In  all  the  cases  hitherto  given  the  male  is  more  strongly  or 
brighter  colored  than  the  female,  and  differs  from  the  young  of 
both  sexes.  But  as  with  some  few  birds  it  is  the  female  which  is 
brighter  colored  than  the  male,  so  with  the  Rhesus  monkey  (Ma- 
cacus  rhesus)  the  female  has  a  large  surface  of  naked  skin  round 
the  tail,  of  a  brilliant  carmine  red,  which,  as  I  was  assured  by 
the  keepers  in  the  Zoological  Gardens,  periodically  becomes  even 
yet  more  vivid,  and  her  face  also  is  pale  red.  On  the  other  hand, 
in  the  adult  male  and  in  the  young  of  both  sexes  (as  I  saw  in 
the  Gardens),  neither  the  naked  skin  at  the  posterior  end  of  the 
body,  nor  the  face,  show  a  trace  of  red.  It  appears,  however, 
from  some  published  accounts,  that  the  male  does  occasionally, 
or  during  certain  seasons,  exhibit  some  traces  of  the  red.  Al- 
though he  is  thus  less  ornamented  than  the  female,  yet  in  the 
larger  size  of  his  body,  larger  canine  teeth,  more  developed  whis- 
kers, more  prominent  superciliary  ridges,  he  follows  the  common 
rule  of  the  male  excelling  the  female. 

I  have  now  given  all  the  cases  known  to  me  of  a  difference  in 
color  between  the  sexes  of  mammals.  Some  of  these  may  be 
the  result  of  variations  confined  to  one  sex  and  transmitted  to 


31  Gervais,  'Hist.  Nat.  des  Mammiferes,'  1854,  p.  103.  Figures  are  given 
of  the  skull  of  the  male.  Also,  Desmarest,  'Mammalogie,'  p.  70.  Geof- 
froy  St.-Hllalre  and  F.  Cuvier,  'Hist.  Nat.  des  Mamm.'  1824,  torn.  i. 


MAMMALS—ORNAMENTAL  COLORS.  535 

the  same  sex,  without  any  good  being  gained,  and  therefore  with- 
out the  aid  of  selection.  We  have  instances  of  this  with  our 
domesticated  animals,  as  in  the  males  of  certain  cats  being  rusty- 
red,  whilst  the  females  are  tortoise-shell  colored.  Analogous 
cases  occur  in  nature:  Mr.  Bartlett  has  seen  many  black  varieties 
of  the  jaguar,  leopard,  vulpine  phalanger,  and  wombat;  and  h.e 
is  certain  that  all,  or  nearly  all  these  animals,  were  males.  On 
the  other  hand,  with  wolves,  foxes,  and  apparently  American 
squirrels,  both  sexes  are  occasionally  born  black.  Hence  it  is 
quite  possible  that  with  some  mammals  a  difference  in  color 
between  the  sexes,  especially  when  this  is  congenital,  may  simply 
be  the  result,  without  the  aid  of  selection,  of  the  occurrence  of 
one  or  more  variations,  which  from  the  first  were  sexually  limited 
in  their  transmission.  Nevertheless  it  is  improbable  that  the 
diversified,  vivid,  and  contrasted  colors  of  certain  quadrupeds, 
for  instance,  of  the  above  monkeys  and  antelopes  can  thus  be 
accounted  for.  We  should  bear  in  mind  that  these  colors  do  not 
appear  in  the  male  at  birth,  but  only  at  or  near  maturity;  and 
that  unlike  ordinary  variations,  they  are  lost  if  the  male  be  emas- 
culated. It  is  on  the  whole  probable  that  the  strongly-marked 
colors  and  other  ornamental  characters  of  male  quadrupeds  are 
beneficial  to  them  in  their  rivalry  with  other  males,  and  have 
consequently  been  acquired  through  sexual  selection.  This  view 
is  strengthened  by  the  differences  in  color  between  the  sexes 
occurring  almost  exclusively,  as  may  be  collected  from  the  pre- 
vious details,  in  those  groups  and  sub-groups  of  mammals,  which 
present  other  and  strongly-marked  secondary  sexual  characters; 
these  being  likewise  due  to  sexual  selection. 

Quadrupeds  manifestly  take  notice  of  color.  Sir  S.  Baker  re- 
peatedly observed  that  the  African  elephant  and  rhinoceros  at- 
tacked white  or  gray  horses  with  special  fury.  I  have  elsewhere 
shown^-  that  half-wild  horses  apparently  prefer  to  pair  with  those 
of  the  same  color,  and  that  herds  of  fallow-deer  of  different  col- 
ors, though  living  together,  have  long  kept  distinct.  It  is  a  more 
significant  fact  that  a  female  zebra  would  not  admit  the  addresses 
of  a  male  ass  until  he  was  painted  so  as  to  resemble  a  zebra,  and 
then,  as  John  Hunter  remarks,  "she  received  him  very  readily. 
"In  this  curious  fact,  we  have  instinct  excited  by  mere  color, 
"which  had  so  strong  an  effect  as  to  get  the  better  of  everything 
"else.  But  the  male  did  not  require  this,  the  female  being  an 
"animal  somewhat  similar  to  himself,  was  sufficient  to  rouse 
"him."=^^ 


82  'The  Variation  of  Animals  and  Plants  under  Domestication,'  1868, 
vol.  ii.  pp.  i02,  103. 

^  'Essays  and  Observations  by  J.  Hunter,'  edited  by  Owen,  1861, 
vol.  i.  p.  194. 


536  THE  DESCENT  OF  MAN. 

In  an  earlier  chapter  we  have  seen  that  the  mental  powers  of 
the  higher  animals  do  not  differ  in  kind,  though  greatly  in  degree, 
from  the  corresponding  powers  of  man,  especially  of  the  lower 
and  barbarous  races;  and  it  would  appear  that  even  their  taste 
for  the  beautiful  is  not  widely  different  from  that  of  the  Quad- 
rumana.  As  the  negro  of  Africa  raises  the  flesh  on  his  face  into 
parallel  ridges  "or  cicatrices,  high  above  the  natural  surface, 
"which  unsightlj'"  deformities,  are  considered  great  personal  at- 
"tractions;"^* — as  negroes  and  savages  in  many  parts  of  the  world 
paint  their  faces  with  red,  blue,  white,  or  black  bars, — so  the  male 
mandrill  of  Africa  appears  to  have  acquired  his  deeply-furrowed 
and  gaudily-colored  face  from  having  been  thus  rendered  attrac- 
tive to  the  female.  No  doubt  it  is  to  us  a  most  grotesque  notion 
that  the  posterior  end  of  the  body  should  be  colored  for  the  sake 
of  ornament  even  more  brilliantly  than  the  face;  but  this  is  not 
more  strange  than  that  the  tails  of  many  birds  should  be  especially 
decorated. 

With  mammals  we  do  not  at  present  possess  any  evidence  that 
the  males  take  pains  to  display  their  charms  before  the  female; 
and  the  elaborate  manner  in  which  this  is  performed  by  male 
birds  and  other  animals,  is  the  strongest  argument  in  favor  of 
the  belief  that  the  females  admire,  or  are  excited  by,  the  orna- 
ments and  colors  displayed  before  them.  There  is,  however,  a 
striking  parallelism  between  mammals  and  birds  in  all  their  sec- 
ondary sexual  characters,  namely  in  their  weapons  for  fighting 
with  rival  males,  in  their  ornamental  appendages,  and  in  their 
colors.  In  both  classes,  when  the  male  differs  from  the  female, 
the  young  of  both  sexes  almost  always  resemble  each  other,  and 
in  a  large  majority  of  cases  resemble  the  adult  female.  In  both 
classes  the  male  assumes  the  characters  proper  to  his  sex  shortly 
before  the  age  of  reproduction;  and  if  emasculated  at  an  early 
period,  loses  them.  In  both  classes  the  change  of  color  is  some- 
times seasonal,  and  the  tints  of  the  naked  parts  sometimes  be- 
come more  vivid  during  the  act  of  courtship.  In  both  classes  the 
male  is  almost  always  more  vividly  or  strongly  colored  than  the 
female,  and  is  ornamented  with  larger  crests  of  hair  or  feathers, 
or  other  such  appendages.  In  a  few  exceptional  cases  the  female 
in  both  classes  is  more  highly  ornamented  than  the  male.  With 
many  mammals,  and  at  least  in  the  case  of  one  bird,  the  male 
is  more  odoriferous  than  the  female.  In  both  classes  the  voice 
of  the  male  is  more  powerful  than  that  of  the  female.  Consider- 
ing this  parallelism  there  can  be  little  doubt  that  the  same  cause, 
whatever  it  may  be,  has  acted  on  mammals  and  birds;  and  the 
result,  as  far  as  ornamental  characters  are  concerned,  may  be  at- 
tributed, as  it  appears  to  me,  to  the  long-continued  preference 

3*  Sir  S,  Baker,   'The  Nile  Tributaries  of  Abyssinia,'  1867. 


MAMMALS— ORNAMENTS.  537 

of  the  individuals  of  one  sex  for  certain  individuals  of  the  opposite 
sex,  combined  with  their  success  in  leaving  a  larger  number  of 
offspring  to  inherit  their  superior  attractions. 

Equal  transmission  of  ornamental  characters  to  hotli  sexes. -^ 
With  many  birds,  ornaments,  which  analogy  leads  us  to  believ© 
were  primarily  acquired  by  the  males,  have  been  transmitted 
equally,  or  almost  equally,  to  both  sexes;  and  we  may  now  in- 
quire how  far  this  view  applies  to  mammals.  With  a  considerable 
number  of  species,  especially  of  the  smaller  kinds,  both  sexes  have 
been  colored,  independently  of  sexual  selection,  for  the  sake  of 
protection;  but  not,  as  far  as  I  can  judge,  in  so  many  cases,  nor 
in  so  striking  a  manner,  as  in  most  of  the  lower  classes.  Audu- 
bon remarks  that  he  often  mistook  the  musk-rat,^^  whilst  sitting 
on  the  banks  of  a  muddy  stream,  for  a  clod  of  earth,  so  complete 
was  the  resemblance.  The  hare  on  her  form  is  a  familiar  instance 
of  concealment  through  color;  yet  this  principle  partly  fails 
in  a  closely-allied  species,  the  rabbit,  for  when  running  to  its 
burrow,  it  is  made  conspicuous  to  the  sportsman,  and  no  doubt 
to  all  beasts  of  prey,  by  its  upturned  white  tail.  No  one  doubts 
that  the  quadrupeds  inhabiting  snow-clad  regions  have  been 
rendered  white  to  protect  them  from  their  enemies,  or  to  favor 
their  stealing  on  their  prey.  In  regions  where  snow  never  lies 
for  long,  a  white  coat  would  be  injurious;  consequently,  species 
of  this  color  are  extremely  rare  in  the  hotter  parts  of  the  world. 
It  deserves  notice  that  many  quadrupeds  inhabiting  moderately 
cold  regions,  although  they  do  not  assume  a  white  winter  dress, 
become  paler  during  this  season;  and  this  apparently  is  the  direct 
result  of  the  conditions  to  which  they  have  long  been  exposed. 
Pallas^*'  states  that  in  Siberia  a  change  of  this  nature  occurs  with 
the  wolf,  two  species  of  Mustela,  the  domestic  horse,  the  Equus 
hemionus,  the  domestic  cow,  two  species  of  antelopes,  the  musk- 
deer,  the  roe,  elk,  and  reindeer.  The  roe,  for  instance,  has  a  red 
summer  and  a  grayish- white  winter  coat;  and  the  latter  may 
perhaps  serve  as  a  protection  to  the  animal  whilst  wandering 
through  the  leafless  thickets,  sprinkled  with  snow  and  hoar-frost. 
If  the  above-named  animals  were  gradually  to  extend  their  range 
into  regions  perpetually  covered  with  snow,  their  pale  winter- 
coats  would  probably  be  rendered  through  natural  selection, 
whiter  and  whiter,  until  they  became  as  white  as  snow. 

Mr.  Reeks  has  given  me  a  curious  instance  of  an  animal  profit- 
ing by  being  peculiarly  colored.     He  raised  from  fifty  to  sixty 

35  Fiber  zibethicus,  Audubon  and  Bachman,  'The  Quadrupeds  of  N. 
America,'  1846,  p.  109. 

38  'Novae  species  Quadrupedum  e  Glirium  ordine,'  1778,  p.  7.  What 
I  have  called  the  roe  is  the  Capreolus  sibiricus  subecaudatus  of  Pallas. 


538  THE  DESCENT  OF  MAN. 

white  and  brown  piebald  rabbits  in  a  large  walled  orchard;  and 
he  had  at  the  same  time  some  similarly  colored  cats  in  his  house. 
Such  cats,  as  I  have  often  noticed,  are  very  conspicuous  during 
day;  but  as  they  used  to  lie  in  watch  during  the  dusk  at  the 
mouths  of  the  burrows,  the  rabbits  apparently  did  not  distinguish 
them  from  their  parti-colored  brethren.  The  result  was  that, 
within  eighteen  months,  every  one  of  these  parti-colored  rabbits 
vv^ere  destroyed;  and  there  was  evidence  that  this  was  effected 
by  the  cats.  Color  seems  to  be  advantageous  to  another  animal, 
the  skunk,  in  a  manner  of  which  we  have  had  many  instances 
in  other  classes.  No  animal  will  voluntarily  attack  one  of  these 
creatures  on  account  of  the  dreadful  odor  which  it  emits  when 
irritated;  but  during  the  dusk  it  would  not  easily  be  recognized 
and  might  be  attacked  by  a  beast  of  prey.  Hence  it  is,  as  Mr. 
Belt  believes,"  that  the  skunk  is  provided  with  a  great  white 
bushy  tail,  which  serves  as  a  conspicuous  warning. 

Although  we  must  admit  that  many  quadrupeds  have  received 
their  present  tints  either  as  a  protection,  or  as  an  aid  in  procuring 
prey,  yet  with  a  host  of  species,  the  colors  are  far  too  conspicuous 
and  too  singularly  arranged  to  allow  us  to  suppose  that  they  serve 
for  these  purposes.    We  may  take  as  an  illustration  certain  ante- 
lopes;   when  we  see  the  square  white  patch  on  the  throat,  the 
white  marks  on  the  fetlocks,  and  the  round  black  spots  on  the 
ears,  all  more  distinct  in  the  male  of  the  Portax  picta  than  in  the 
female; — when  we  see  that  the  colors  are  more  vivid,  that  the 
narrow  white  lines  on  the  flank  and  the  broad  white  bar  on  the 
shoulder  are  more  distinct  in  the  male  Oreas  derbyanus  than  in 
the  female; — when  we  see  a  similar  difference  between  the  sexes 
of  the  curiously-ornamented  Tragelaphus  scriptus  (fig.  70),— we 
cannot  believe  that  differences  of  this  kind  are  of  any  service  to 
either  sex  in  their  daily  habits  of  life.     It  seems  a  much  more 
probable  conclusion  that  the  various  marks  were  first  acquired 
by  the  males  and  their  colors  intensified  through  sexual  selection, 
and  then  partially  transferred  to  the  females.     If  this  view  be 
admitted,  there  can  be  little  doubt  that  the  equally  singular  colors 
and  marks  of  many  other  antelopes,  though  common  to  both 
sexes,  have  been  gained  and  transmitted  in  a  like  manner.    Both 
sexes,  for  instance,  of  the  koodoo   (Strepsiceros  kudu)   (fig.  64) 
have  narrow  white  vertical  lines  on  their  hind  flanks,  and  an 
elegant  angular  white  mark  on  their  foreheads.     Both  sexes  in 
the  genus  Damalis  are  very  oddly  colored;    in  D.  pygarga  the 
back  and  neck  are  purplish-red,  shading  on  the  flanks  into  black; 
and  these  colors  are  abruptly  separated  from  the  white  belly  and 
from  a  large  white  space  on  the  buttocks;    the  head  is  still  more 


8'  'The  Naturalist  in  Nicaragua,'  p.  249. 


MAMMALS— SPOTS  AND  STRIPES. 


§39 


Oddly  colored,  a  large  oblong  -v^hite  mask,  narrowly-edged  with 
black,  covers  the  face  up  to  the  eyes  (fig.  71) ;  there  are  three 
white  stripes  on  the  forehead,  and  the  ears  are  marked  with 
white.  The  fawns  of  this  species  are  of  a  uniform  pale  yellowish- 
brown.  In  Damalis  albifrons  the  coloring  of  the  head  differs  from 
that  in  the  last  species  in  a  single  white  stripe  replacing  the  three 
stripes,  and  in  the  ears  being  almost  wholly  white.^   After  having 


Fig.  70.    Tragelaphus  scriptus,  male  (from  the  Knowsley  Menagerie). 

Studied  to  the  best  of  my  ability  the  sexual  differences  of  animals 
belonging  to  all  classes,  I  cannot  avoid  the  conclusion  that  the 
curiously-arranged  colors  of  many  antelopes,  though  common  to 
both  sexes,  are  the  result  of  sexual  selection  primarily  applied  to 
the  male. 


38  See  the  fine  plates  in  A.  Smith's  ^Zoology  of  S.  Africa,*  and   Dr. 
Gray's  'Gleanings  from  the  Menagerie  of  KnowS'ley.* 


540 


THE  DESCENT  OF  MAN. 


The  same  conclusion  may  perhaps  be  extended  to  the  tiger,  one 
of  the  most  beautiful  animals  in  the  world,  the  sexes  of  which 
cannot  be  distinguished  by  color,  even  by  the  dealers  in  wild 
beasts.  Mr.  Wallace  believes^^  that  the  striped  coat  of  the  tiger 
**so  assimilates  with  the  vertical  stems  of  the  bamboo,  as  to 
"assist  greatly  in  concealing  him  from  his  approaching  prey." 
But  this  view  does  not  appear  to  me  satisfactory.    We  have  some 


Pig.  71.    Damalis  pygarg-a,   male  (from  the  Knowsley  Menagerie). 

slight  evidence  that  his  beauty  may  be  due  to  sexual  selection, 
for  in  two  species  of  Felis  the  analogous  marks  and  colors  are 
rather  brighter  in  the  male  than  in  the  female.  The  zebra  is 
conspicuously  striped,  and  stripes  cannot  afford  any  protection 
on  the  open  plains  of  South  Africa.  BurchelP°  in  describing  a 
herd  says,  "their  sleek  ribs  glistened  in  the  sun,  and  the  bright- 
"ness  and  regularity  of  their  striped  coats  presented  a  picture  of 
"extraordinary  beauty,  in  which  probably  they  are  not  surpassed 


a"  'Westminster  Review,'  July  1,  1867,  p.  5. 

40  'Travels  in  South  Africa,'  1824.  vol.  ii.  p.  315. 


MAMMALS— SPOTS  AND  STRIPES.  541 

"by  any  other  quadruped."  But  as  throughout  the  whole  group 
of  the  Equidse  the  sexes  are  identical  in  color,  we  have  here  no 
evidence  of  sexual  selection.  Nevertheless  he  who  attributes  the 
white  and  dark  vertical  stripes  on  the  flanks  of  various  antelopes 
to  this  process,  will  probably  extend  the  same  view  to  the  Royal 
Tiger  and  beautiful  Zebra. 

We  have  seen  in  a  former  chapter  that  when  young  animals 
belonging  to  any  class  follow  nearly  the  same  habits  of  life  as 
their  parents,  and  yet  are  colored  in  a  different  manner,  it  may 
be  inferred  that  they  have  retained  the  coloring  of  some  ancient 
and  extinct  progenitor.  In  the  family  of  pigs,  and  in  the  tapirs, 
the  young  are  marked  with  longitudinal  stripes,  and  thus  differ 
from  all  the  existing  adult  species  in  these  two  groups.  With 
many  kinds  of  deer  the  young  are  marked  with  elegant  white 
spots,  of  which  their  parents  exhibit  not  a  trace.  A  graduated 
series  can  be  followed  from  the  axis  deer,  both  sexes  of  which  at 
all  ages  and  during  ail  seasons  are  beautifully  spotted  (the  male 
being  rather  more  strongly  colored  than  the  female),  to  species 
in  which  neither  the  old  nor  the  young  are  spotted.  I  will  specify 
some  of  the  steps  in  this  series.  The  Mantchurian  deer  (Cervus 
mantchuricus)  is  spotted  during  the  whole  year,  but,  as  I  have 
seen  in  the  Zoological  Gardens,  the  spots  are  much  plainer  during 
the  summer,  when  the  general  color  of  the  coat  is  lighter,  than 
during  the  winter,  when  the  general  color  is  darker  and  the  horns 
are  fully  developed.  In  the  hog-deer  (Hyelaphus  porcinus)  the 
spots  are  extremely  conspicuous  during  the  summer  when  the 
coat  is  reddish-brown,  but  quite  disappear  during  the  winter 
when  the  coat  is  brown.^  In  both  these  species  the  young  are 
spotted.  In  the  Virginian  deer  the  young  are  likewise  spotted, 
and  about  five  per  cent,  of  the  adult  animals  living  in  Judge 
Caton's  park,  as  I  am  informed  by  him,  temporarily  exhibit  at  the 
period  when  the  red  summer  coat  is  being  replaced  by  the  bluish 
winter  coat,  a  row  of  spots  on  each  flank,  which  are  always  the 
same  in  number,  though  very  variable  in  distinctness.  From  this 
condition  there  is  but  a  very  small  step  to  the  complete  absence 
of  spots  in  the  adults  at  all  seasons;  and,  lastly,  to  their  absence 
at  all  ages  and  seasons,  as  occurs  with  certain  species.  From  the 
existence  of  this  perfect  series,  and  more  especially  from  the 
fawns  of  so  many  species  being  spotted,  we  may  conclude  that  the 
now  living  members  of  the  deer  family  are  the  descendants  of 
some  ancient  species  which,  like  the  axis  deer,  was  spotted  at  all 
ages  and  seasons.    A  still  more  ancient  progenitor  probably  some- 

"  Dr.  Gray,  'Gleanings  from  the  Menagerie  of  Knowsley,'  p.  64.  Mr. 
Blyth,  in  speaking  ('Land  and  Water,'  1869,  p.  42)  of  the  hog-deer  of 
Ceylon,  says  it  is  more  brightly  spotted  with  white  than  the  common 
hog-deer,  at  the  season  when  it  renews  its  horns. 


542  '  THE  DESCENT  OF  MAN. 

what  resembled  the  Hyomosclius  aquaticus — for  this  animal  is 
spotted,  and  the  hornless  males  have  large  exserted  canine  teeth, 
of  which  some  few  true  deer  still  retain  rudiments.  Hyomoschus, 
also,  offers  one  of  those  interesting  cases  of  a  form  linking  to- 
gether two  groups,  for  it  is  intermediate  in  certain  osteological 
characters  between  the  pachyderms  and  ruminants,  which  were 
formerly  thought  to  be  quite  distinct.*^ 

A  curious  difficulty  here  arises.  If  we  admit  that  colored  spots 
and  stripes  were  first  acquired  as  ornaments,  how  comes  it  that 
so  many  existing  deer,  the  descendants  of  an  aboriginally  spotted 
animal,  and  all  the  species  of  pigs  and  tapirs,  the  descendants  of 
an  aboriginally  striped  animal,  have  lost  in  their  adult  state  their 
former  ornaments?  I  cannot  satisfactorily  answer  this  question. 
We  may  feel  almost  sure  that  the  spots  and  stripes  disappeared 
at  or  near  maturity  in  the  progenitors  of  our  existing  species,  so 
that  they  were  still  retained  by  the  young;  and,  owing  to  the  law 
of  inheritance  at  corresponding  ages,  were  transmitted  to  the 
young  of  all  succeeding  generations.  It  may  have  been  a  great 
advantage  to  the  lion  and  puma,  from  the  open  nature  of  their 
usual  haunts,  to  have  lost  their  stripes,  and  to  have  been  thus 
rendered  less  conspicuous  to  their  prey;  and  if  the  successive 
variations,  by  which  this  end  was  gained,  occurred  rather  late  in 
life,  the  young  would  have  retained  their  stripes,  as  is  now  the 
case.  As  to  deer,  pigs,  and  tapirs,  Fritz  Miiller  has  suggested 
to  me  that  these  animals,  by  the  removal  of  their  spots  or  stripes 
through  natural  selection,  would  have  been  less  easily  seen  by 
their  enemies;  and  that  they  w^ould  have  especially  required  this 
protection,  as  soon  as  the  carnivora  increased  in  size  and  number 
during  the  tertiary  periods.  This  may  be  the  true  explanation, 
but  it  is  rather  strange  that  the  young  should  not  have  been  thus 
protected,  and  still  more  so  that  the  adults  of  some  species  should 
have  retained  their  spots,  either  partially  or  completely,  during 
part  of  the  year.  We  know  that,  when  the  domestic  ass  varies 
and  becomes  reddish-brown,  gray,  or  black,  the  stripes  on  the 
shoulders  and  even  on  the  spine  frequently  disappear,  though  we 
cannot  explain  the  cause.  Very  few  horses,  except  dun-colored 
kinds,  have  stripes  on  any  part  of  their  bodies,  yet  we  have  good 
reason  to  believe  that  the  aboriginal  horse  was  striped  on  the 
legs  and  spine,  and  probably  on  the  shoulders.^  Hence  the  dis- 
appearance of  the  spots  and  stripes  in  our  adult  existing  deer, 
pigs,  and  tapirs,  may  be  due  to  a  change  in  the  general  color  of 
their  coats;    but  whether  this  change  was  effected  through  sexual 


*2  Falconer  and  Cautley,  'Proc.  Geolog.  Soc'  1843;  and  Falconer's 
'Pal.  Memoirs/  vol.  i.  p.  196. 

^  'The  Variation  of  Animals  and  Plants  under  Domestication,'  1868, 
vol.  i.  p.  61-64. 


MAMMALS— BEAUTY  OF  THE  QUADRUMANA. 


;43 


or  natural  selection,  or  was  due  to  the  direct  action  of  the  condi- 
tions of  life,  or  to  some  other  unknown  cause,  it  is  impossible 
to  decide.  An  observation  made  by  Mr.  Sclater  well  illustrates 
our  ignorance  of  the  laws  which  regulate  the  appearance  and 
disappearance  of  stripes;  the  species  of  Asinus  which  inhabit 
the  Asiatic  continent  are  destitute  of  stripes,  not  having  even 


Fig.  72.  Head  of  Semnopithecus  rubicundus.  This  and  the  following 
fig-ures  (from  Prof.  Gervais)  are  g-iven  to  show  the  odd  arrange- 
ment and  development  of  the  hair  on  the  head. 

the  cross  shoulder-stripe,  whilst  those  which  inhabit  Africa  are 
conspicuously  striped,  with  the  partial  exception  of  A.  tseniopus, 
which  has  only  the  cross  shoulder-stripe  and  generally  some 
faint  bars  on  the  legs;  and  this  species  inhabits  the  almost  inter- 
mediate region  of  Upper  Egypt  and  Abyssinia.^ 

Quadrumana. — Before  we  conclude,  it  will   be  well  to  add  a 
few  remarks  on  the  ornaments  of  monkeys.    In  most  of  the  spe- 


^  'Proc.  Zool.  Soc'  1862,  p.  164.    See,   also,   Dr.   Hartmann,    'Ann.   d. 
Landw.'    Bd.  xliii.  s.  222. 


514 


THE  DESCENT  OF  MAN. 


cies  the  sexes  resemble  each  other  in  color,  but  in  some,  as  we 
have  seen,  the  males  differ  from  the  females,  especially  in  the 
color  of  the  naked  parts  of  the  skin,  in  the  development  of  the 
beard,  whiskers,  and  mane.  Many  species  are  colored  either  in 
so  extraordinary  or  so  beautiful  a  manner,  and  are  furnished 
with  such  curious  and  elegant  crests  of  hair,  that  we  can  hardly 
avoid  looking  at  these  characters  as  having  been  gained  for  the 


Fig.  73.    Head  of  Semnopithecus 
comatus. 


Fig-.  74.    Head  of  Cebus  capucinus. 


Fig.  75.    Head  of  Ateles  marginatus. 


Fig.  76.    Head  of  Cebus 
vellerosus. 


sake  of  ornament.  The  accompanying  figures  (figs.  72  to  76) 
serve  to  show  the  arrangement  of  the  hair  on  the  face  and  head 
in  several  species.  It  is  scarcely  conceivable  that  these  crests  of 
hair,  and  the  strongly  contrasted  colors  of  the  fur  and  skin,  can 
be  the  result  of  mere  variability  without  the  aid  of  selection; 
and  it  is  inconceivable  that  they  can  be  of  use  in  any  ordinary 
way  to  these  animals.  If  so,  they  have  probably  been  gained 
through  sexual  selection,  though  transmitted  equally,  or  almost 


BEAUTY  OF  THE  QUADRUMANA. 


545 


equally,  to  both  sexes.  With  many  of  the  Quadrumana,  we  have 
additional  evidence  of  the  action  of  sexual  selection  in  the  greater 
size  and  strength  of  the  males,  and  in  the  greater  development  of 
their  canine  teeth,  in  comparison  with  the  females. 


Fig.  77,    Cercopithecus  petaurista  (from  Brehm). 
A  few  instances  will  suffice  of  the  strange  manner  in  which 
both  sexes  of  some  species  are  colored,  and  of  the  beauty  of  others. 
The  face  of  the  Cercopithecus  petaurista  (fig.  77)   is  black,  the 
36 


546  THE  DESCENT  OP  MAN. 

whiskers  and  beard  being  white,  with  a  defined,  round,  white 
spot  on  the  nose,  covered  with  short  white  hair,  which  gives  to 
the  animal  an  almost  ludicrous  aspect.  The  Semnopithecus  fron- 
tatus  likewise  has  a  blackish  face  with  a  long  black  beard,  and 
a  large  naked  spot  on  the  forehead  of  a  bluish-white  color.  The 
face  of  the  Macacus  lasiotus  is  dirty  flesh-colored,  with  a  defined 
red  spot  on  each  cheek.  The  appearance  of  Cercocebus  a3thiops 
is  grotesque,  with  its  black  face,  white  whiskers  and  collar,  chest- 
nut head,  and  a  large  naked  white  spot  over  each  eyelid.  In  very 
many  species,  the  beard,  v/hiskers,  and  crests  of  hair  round  the 
face  are  of  a  different  color  from  the  rest  of  the  head,  and  when 
different,  are  always  of  a  lighter  tint,*^  being  often  pure  white, 
sometimes  bright  yellow,  or  reddish.  The  whole  face  of  the  South 
American  Brachyurus  calvus  is  of  a  "glowing  scarlet  hue;"  but 
this  color  does  not  appear  until  the  animal  is  nearly  mature.*® 
The  naked  skin  of  the  face  differs  wonderfully  in  color  in  the 
various  species.  It  is  often  brown  or  flesh-color,  with  parts  per- 
fectly white,  and  often  as  black  as  that  of  the  most  sooty  negro. 
In  the  Brachyurus  the  scarlet  tint  is  brighter  than  that  of  the 
most  blushing  Caucasian  damsel.  It  is  sometimes  more  distinctly 
orange  than  in  any  Mongolian,  and  in  several  species  it  is  blue, 
passing  into  violet  or  gray.  In  all  the  species  known  to  Mr.  Bart- 
lett,  in  which  the  adults  of  both  sexes  have  strongly-colored 
faces,  the  colors  are  dull  or  absent  during  early  youth.  This  like^ 
wise  holds  good  with  the  mandrill  and  Rhesus,  in  which  the  face 
and  the  posterior  parts  of  the  body  are  brilliantly  colored  in  one 
sex  alone.  In  these  latter  cases  we  have  reason  to  believe  that 
the  colors  were  acquired  through  sexual  selection;  and  we  are 
naturally  led  to  extend  the  same  view  to  the  foregoing  species, 
though  both  sexes  when  adult  have  their  faces  colored  in  the 
same  manner. 

Although  many  kinds  of  monkeys  are  far  from  beautiful  accord- 
ing to  our  taste,  other  species  are  universally  admired  for  their 
elegant  appearance  and  bright  colors.  The  Semnopithecus  ne- 
mseus,  though  peculiarly  colored,  is  described  as  extremely  pretty; 
the  orange-tinted  face  is  surrounded  by  long  whiskers  of  glossy 
whiteness,  with  a  line  of  chestnut-red  over  the  eyebrows;  the 
fur  on  the  back  is  of  a  delicate  gray,  with  a  square  patch  on  the 
loins,  the  tail  and  the  fore-arms  being  of  a  pure  white;  a  gorget 
of  chestnut  surmounts  the  chest;  the  thighs  are  black,  with  the 
legs  chestnut  red.  I  will  mention  only  two  other  monkeys  for 
their  beauty;    and  I  have  selected  these  as  presenting  slight  sex- 


*5 1  observed  this  fact  in  the  Zoological  Gardens;  and  many  cases 
may  be  seen  in  the  colored  plates  in  Geoffrey  St.-Hilaire  and  F.  Cu- 
yier,  'Hist.  Nat,  des  Mammiferes,'  tom.  i.  1824. 

*8  Bates,  'The  Naturalist  on  the  Amazons,'  1863,  vol.  ii.  p.  310. 


BEAUTY  OF  THE  QUADRUMANA. 


547 


iial  differences  in  color,  whicli  renders  it  in  some  degree  probable 
that  both  sexes  owe  their  elegant  appearance  to  sexual  selection. 
In  the  moustache-monkey  (Cercopithecus  cephus)  the  general 
color  of  the  fur  is  mottled-greenish  with  the  throat  white;  in 
the  male  the  end  of  the  tail  is  chestnut,  but  the  face  is  the  most 
ornamented  part,  the  skin  being  chiefly  bluish-gray,  shading  into 


Fig.  78,    Cercopithecus  diana  (from  Brehm). 

a  blackish  tint  beneath  the  eyes,  with  the  upper  lip  of  a  delicate 
blue,  clothed  on  the  lower  edge  with  a  thin  black  moustache;  the 
whiskers  are  orange-colored,  with  the  upper  part  black,  forming 
a  band  which  extends  backv/ards  to  the  ears,  the  latter  being 
clothed  with  whitish  hairs.  In  the  Zoological  Society's  Gardens 
I  have  often  overheard  visitors  admiring  the  beauty  of  another 
monkey,  deservedly  called  Cercopithecus  diana  (fig.  78);  the 
general  color  of  the  fur  is  gray;   the  chest  and  inner  surface  of  the 


G48  THE  DESCENT  OF  MAN. 

forelegs  are  white;  a  large  triangular  defined  space  on  the  hinder 
part  of  the  back  is  rich  chestnut;  in  the  male  the  inner  sides 
of  the  thighs  and  the  abdomen  are  delicate  fawn-colored,  and 
the  top  of  the  head  is  black;  the  face  and  ears  are  intensely 
black,  contrasting  finely  with  a  white  transverse  crest  over  the 
eyebrows  and  a  long  white  peaked  beard,  of  which  the  basal  por- 
tion is  black/^ 

In  these  and  many  other  monkeys,  the  beauty  and  singular 
arrangement  of  their  colors,  and  still  more  the  diversified  and 
elegant  arrangement  of  the  crests  and  tufts  of  hair  on  their 
heads,  force  the  conviction  on  my  mind  that  these  characters 
have  been  acquired  through  sexual  selection  exclusively  as  orna- 
ments. 

Summm^y,  — The  law  of  battle  for  the  possession  of  the  female 
appears  to  prevail  throughout  the  whole  great  class  of  mammals. 
Most  naturalists  will  admit  that  the  greater  size,  strength,  cour- 
age, and  pugnacity  of  the  male,  his  special  weapons  of  offense, 
as  well  as  his  special  means  of  defense,  have  been  acquired  or 
modified  through  that  form  of  selection  which  I  have  called 
sexual.  This  does  not  depend  on  any  superiority  in  the  general 
struggle  for  life,  but  on  certain  individuals  of  one  sex,  generally 
the  male,  being  successful  in  conquering  other  males,  and  leaving 
a  larger  number  of  offspring  to  inherit  their  superiority  than  do 
the  less  successful  males. 

There  is  another  and  more  peaceful  kind  of  contest,  in  which 
the  males  endeavor  to  excite  or  allure  the  females  by  various 
charms.  This  is  probably  carried  on  in  some  cases  by  the  power- 
ful odors  emitted  by  the  males  during  the  breeding-season;  the 
odoriferous  glands  having  been  acquired  through  sexual  selec- 
tion. Whether  the  same  view  can  be  extended  to  the  voice  is 
doubtful,  for  the  vocal  organs  of  the  males  must  have  been 
strengthened  by  use  during  maturity,  under  the  powerful  excite- 
ments of  love,  jealousy  or  rage,  and  will  consequently  have  been 
transmitted  to  the  same  sex.  Various  crests,  tufts,  and  mantles 
of  hair,  which  are  either  confined  to  the  male,  or  are  more  de- 
veloped in  this  sex  than  in  the  female,  seem  in  most  cases  to  be 
merely  ornamental,  though  they  sometimes  serve  as  a  defense 
against  rival  males.  There  is  even  reason  to  suspect  that  the 
branching  horns  of  stags,  and  the  elegant  horns  of  certain  ante- 
lopes, though  properly  serving  as  weapons  of  offense  or  defense, 
have  been  partly  modified  for  ornament. 


^"^  I  have  seen  most  of  the  above  monkeys  in  the  Zoological  Society's 
Gardens.  The  description  of  the  Semnopithecus  nemaeus  is  taken  from 
Mr.  W.  C.  Martin's  'Nat.  Hist,  of  Mammalia,'  1841,  p.  460;  see.  also, 
pp.  475,  523. 


MAMMALS— SUMMARY.  549 

When  the  male  differs  in  color  from  the  female,  he  generally 
exhibits  darker  and  more  strongly-contrasted  tints.  We  do  not 
in  this  class  meet  with  the  splendid  red,  blue,  yellow,  and  green 
tints,  so  common  with  male  birds  and  many  other  animals.  The 
naked  parts,  however,  of  certain  Quadrumana  must  be  excepted; 
for  such  parts,  often  oddly  situated,  are  brilliantly  colored  in 
some  species.  The  colors  of  the  male  in  other  cases  may  be 
due  to  simple  variation,  without  the  aid  of  selection.  But  when 
the  colors  are  diversified  and  strongly  pronounced,  when  they 
are  not  developed  until  near  maturity,  and  when  they  are  lost 
after  emasculation,  we  can  hardly  avoid  the  conclusion  that  they 
have  been  acquired  through  sexual  selection,  for  the  sake  of  orna- 
ment, and  have  been  transmitted  exclusively,  or  almost  exclusive- 
ly, to  the  same  sex.  When  both  sexes  are  colored  in  the  same 
manner,  and  the  colors  are  conspicuous  or  curiously  arranged, 
without  being  of  the  least  appa,rent  use  as  a  protection,  and  espe- 
cially when  they  are  associated  with  various  other  ornamental 
appendages,  we  are  led  by  analogy  to  the  same  conclusion,  name- 
ly, that  they  have  been  acquired  through  sexual  selection,  al- 
though transmitted  to  both  sexes.  That  conspicuous  and  diversi- 
fied colors,  whether  confined  to  the  males  or  common  to  both 
sexes,  are  as  a  general  rule  associated  in  the  same  groups  and 
sub-groups  with  other  secondary  sexual  characters  serving  for 
war  or  for  ornament,  will  be  found  to  hold  good,  if  Vv'-e  look  back 
to  the  various  cases  given  in  this  and  the  last  chapter. 

The  law  of  the  equal  transmission  of  characters  to  both  sexes, 
as  far  as  color  and  other  ornaments  are  concerned,  has  prevailed 
far  more  extensively  with  mammals  than  with  birds;  but  weap- 
ons, such  as  horns  and  tusks,  have  often  been  transmitted  either 
exclusively  or  much  more  perfectly  to  the  males  than  to  the  fe- 
males. This  is  surprising,  for,  as  the  males  generally  use  their 
v/eapons  for  defense  against  enemies  of  all  kinds,  their  weapons 
would  have  been  of  service  to  the  females.  As  far  as  we  can  see, 
their  absence  in  this  sex  can  be  accounted  for  only  by  the  form 
of  inheritance  which  has  prevailed.  Finally,  with  quadrupeds  the 
contest  between  the  individuals  of  the  same  sex,  whether  peace- 
ful or  bloody,  has,  with  the  rarest  exceptions,  been  confined  to 
the  males;  so  that  the  latter  have  been  modified  through  sexual 
selection,  far  more  commonly  than  the  females,  either  for  fight- 
ing with  each  other  or  for  alluring  the  opposite  sex. 


MAN— SEXUAL.  DIFFERENCES.  551 


PART  III. 


SEXUAL  SELECTION  IN  RELATION  TO  MAN, 

AND    CONCLUSION. 


CHAPTER   XIX. 
SECONDARY    SEXUAL    CHARACTERS    OF    MAN. 

Differences  between  man  and  woman — Causes  of  such  differences  and 
of  certain  characters  common  to  both  sexes — Law  of  battle — Dif- 
ferences in  mental  powers,  and  voice — On  the  influence  of  beauty 
in  determining"  the  marriages  of  mankind — Attention  paid  by  sav- 
ages to  ornaments— Their  ideas  of  beauty  in  woman — The  tendency 
to  exag-g-erate  each  natural  peculiarity. 

With  mankind  the  differences  between  the  sexes  are  greater 
than  in  most  of  the  Quadrumana,  but  not  so  great  as  in  some, 
for  instance,  the  mandrill.  Man  on  an  average  is  considerably 
taller,  heavier,  and  stronger  than  woman,  with  squarer  shoulders 
and  more  plainly-pronounced  muscles.  Owing  to  the  relation 
which  exists  between  muscular  development  and  the  projection  of 
the  brows,^  the  superciliary  ridge  is  generally  more  marked  in 
man  than  in  woman.  His  body,  and  especially  his  face,  is  more 
hairy,  and  his  voice  has  a  different  and  more  powerful  tone.  In 
certain  races  the  women  are  said  to  differ  slightly  in  tint  from 
the  men.  For  instance,  Schweinfurth,  in  speaking  of  a  negress 
belonging  to  the  Monbuttoos,  who  inhabit  the  interior  of  Africa  a 
few  degrees  north  of  the  Equator,  says,  "Like  all  her  race,  she  had 
"a  skin  several  shades  lighter  than  her  husband's,  being  some- 
"thing  of  the  color  of  half -roasted  coffee."-    As  the  women  labor 


1  Schaaffhausen,    translation  in  'Anthropological  Review,'   Oct.  1868, 
pp.  419,  420,  427. 

2  'The  Heart  of  Africa,'  English  Transl.  1873,  vol.  i.  p.  544. 


V 


552  THE   DESCENT  OF  MAN. 

in  the  fields  and  are  quite  unclothed,  it  is  not  likely  that  they 
differ  in  color  from  the  men  owing  to  less  exposure  to  the  weath- 
er. European  women  are  perhaps  the  brighter  colored  of  the 
two  sexes,  as  may  he  seen  when  both  have  been  equally  ex- 
posed. 
^  Man  is  more  courageous,  pugnacious  and  energetic  than  woman, 
and  has  a  more  inventive  genius.  His  brain  is  absolutely  larger, 
but  whether  or  not  proportionately  to  his  larger  body,  has  not,  I 
believe,  been  fully  ascertained.  In  woman  the  face  is  rounder; 
the  jaws  and  the  base  of  the  skull  smaller;  the  outlines  of  the 
body  rounder,  in  parts  more  prominent;  and  her  pelvis  is  broader 
than  in  man;^  but  this  latter  character  may  perhaps  be  considered 
rather  as  a  primary  than  a  secondary  sexual  character.  She 
comes  to  maturity  at  an  earlier  age  than  man. 

As  with  animals  of  all  classes,  so  with  man,  the  distinctive 
characters  of  the  male  sex  are  not  fully  developed  until  he  is 
nearly  mature;  and  if  emasculated  they  never  appear.  The 
beard,  for  instance,  is  a  secondary  sexual  character,  and  male 
children  are  beardless,  though  at  an  early  age  they  have  abundant 
hair  on  the  head.  It  is  probably  due  to  the  rather  late  appearance 
in  life  of  the  successive  variations  v/hereby  man  has  acquired  liis 
masculine  characters,  that  they  are  transmitted  to  the  male  sex 
alone.  Male  and  female  children  resemble  each  other  closely, 
like  the  young  of  so  many  other  animals  in  which  the  adult  sexes 
differ  widely;  they  likevvrise  resemble  the  mature  female  much 
more  closely  than  the  mature  male.  The  female,  however,  ulti- 
mately assumes  certain  distinctive  characters,  and  in  the  forma- 
tion of  her  skull,  is  said  to  be  intermediate  between  the  child 
and  the  man.*  Again,  as  the  young  of  closely  allied  though  dis- 
tinct species  do  not  differ  nearly  so  much  from  each  other  as  do 
the  adults,  so  it  is  with  the  children  of  the  different  races  of  man. 
Some  have  even  maintained  that  race-differences  cannot  be  de- 
tected in  the  infantile  skull.^  In  regard  to  color,  the  new-born 
negro  child  is  reddish  nut-brov/n,  which  soon  becomes  slaty-gray; 
the  black  color  being  fully  developed  within  a  year  in  the  Soudan, 
but  not  until  three  years  in  Egypt.  The  eyes  of  the  negro  are  at 
first  blue,  and  the  hair  chestnut-brown  rather  than  black,  being 
curled  only  a,t  the  ends.  The  children  of  the  Australians  imme- 
diately after  birth  are  yellowish-brown,  and  become  dark  at  a 
later  age.    Those  of  the  Guaranys  of  Paraguay  are  whitish-yellow, 

3  Ecker,  translation  in  'Anthropological  Review,'  Oct.  1868,  pp.  351- 
35(5.  The  comparison  of  the  form  of  the  skull  in  men  and  women  has 
been  followed  out  with  much  care  by  Welcker. 

^  Ecker  and  Welcker,  ibid.  p.  352,  355;  Vogt,  'Lectures  on  Man,'  En^. 
translat.  p.  81. 

^  Schaaffhausen,  'Anthropolog.  Review,'  ibid.  p.  429. 


MAN— SEXUAL   DIFFERENCES.  553 

but  they  acquire  in  the  course  of  a  few  weeks  the  yellowish-brown 
tint  of  their  parents.  Similar  observations  have  been  made  in 
other  parts  of  America.® 

I  have  specified  the  foregoing  differences  between  the  male  and 
female  sex  in  mankind,  because  they  are  curiously  like  those  of 
the  Quadrumana.  With  these  animals  the  female  is  mature  at 
an  earlier  age  than  the  male;  at  least  this  is  certainly  the  case  in 
the  Cebus  azarse=^  The  males  of  most  species  are  larger  and 
stronger  than  the  females,  of  which  fact  the  gorilla  affords  a  well- 
known  instance.  Even  in  so  trifling  a  character  as  the  greater 
prominence  of  the  superciliary  ridge,  the  males  of  certain  monkeys 
differ  from  the  females,®  and  agree  in  this  respect  with  mankind. 
In  the  gorilla  and  certain  other  monkeys,  the  cranium  of  the 
adult  male  presents  a  strongly-marked  sagittal  crest,  which  is 
absent  in  the  female;  and  Ecker  found  a  trace  of  a  similar  dif- 
ference between  the  two  sexes  in  the  Australians.^  With  monkeys 
when  there  is  any  difference  in  the  voice,  that  of  the  male  is  the 
more  powerful.  We  have  seen  that  certain  male  monkeys  have 
a  well-developed  beard,  which  is  quite  deficient,  or  much  less  de- 
veloped in  the  female.  No  instance  is  known  of  the  beard,  whis- 
kers, or  moustache  being  larger  in  the  female  than  in  the  male 
monkey.  Even  in  the  color  of  the  beard  there  is  a  curious  paral- 
lelism between  man  and  the  Quadrumana,  for  with  man  when 
the  beard  differs  in  color  from  the  hair  of  the  head,  as  is  commonly 
the  case,  it  is,  I  believe,  almost  always  of  a  lighter  tint,  being 
often  reddish.  I  have  repeatedly  observed  this  fact  in  England; 
but  two  gentlemen  have  lately  written  to  me,  saying  that  they 
form  an  exception  to  the  rule.  One  of  these  gentlemen  accounts 
for  the  fact  by  the  wide  difference  in  color  of  the  hair  on  the 
paternal  and  maternal  sides,  of  his  family.  Both  had  been  long 
aware  of  this  peculiarity  (one  of  them  having  often  been  accused 
of  dyeing  his  beard),  and  had  been  thus  led  to  observe  other  men, 
and  were  convinced  that  the  exceptions  were  very  rare.  Dr. 
Hooker  attended  to  this  little  point  for  me  in  Russia,  and  found 
no  exception  to  the  rule.  In  Calcutta,  Mr.  J.  Scott,  of  the  Botanic 
Gardens,  was  so  kind  as  to  observe  the  many  races  of  men  to  be 

« Pruner-Bey,  on  negro  infants  as  quoted  by  Vog-t,  'Lectures  on 
Man,'  Eng-.  translat.  1864,  p.  189:  for  further  facts  on  negro  infants, 
as  quoted  from  Winterbottom  and  Camper,  see  Lawrence,  'Lectures 
on  Physiology,'  «S;c.,  1822,  p.  451.  For  the  infants  of  the  Guaranys,  see 
Rengger,  'Saugethiere,'  «S:c.,  s.  3.  See,  also,  Godron,  'De  I'Espece,' 
torn.  ii.  1859,  p.  253.  For  the  Australians,  Waitz,  'Introduct.  to  Anthro- 
polog-y,'  Eng.  translat.  1863,  p.  99. 

'  Rengger,  'Saugethiere,'  &c.,  1830,  s.  49. 

8  As  in  Macacus  cynomolgus  (Desmarest,  'Mammalogie,'  p.  65),  and 
in  Hylobates  agilis  (Geoffrey  St.-Hilaire  and  F.  Cuvier,  'Hist.  Nat. 
des  Mamm.'  1824,  torn.  i.  p.  2). 

»  'Anthropological  Review,'  Oct.  1868,  p.  353. 


554  THE  DESCENT  OF  MAN. 

seen  there,  as  well  as  in  some  other  parts  of  India,  namely  two 
races  in  Sikhim,  the  Bhoteas,  Hindoos,  Burmese,  and  Chinese, 
most  of  which  races  have  very  little  hair  on  the  face;  and  he 
always  found  that  when  there  was  any  difference  in  color  between 
the  hair  of  the  head  and  the  beard,  the  latter  was  invariably 
lighter.  Now  with  monkeys,  as  has  already  been  stated,  the 
beard  frequently  differs  strikingly  in  color  from  the  hair  of  the 
head,  and  in  such  cases  it  is  always  of  a  lighter  hue,  being  often 
pure  white,  sometimes  yellow  or  reddish.^** 

In  regard  to  the  general  hairiness  of  the  body,  the  women  in 
all  races  are  less  hairy  than  the  men;  and  in  some  few  Quad- 
rumana  the  under  side  of  the  body  of  the  female  is  less  hairy 
that  that  of  the  male."  Lastly,  male  monkeys,  like  men,  are 
bolder  and  fiercer  than  the  females.  They  lead  the  troop,  and 
when  there  is  danger,  come  to  the  front.  We  thus  see  how  close 
is  the  parallelism  between  the  sexual  differences  of  man  and  the 
Quadrumana.  With  some  few  species,  however,  as  with  certain 
baboons,  the  orang  and  the  gorilla,  there  is  a  considerably  greater 
difference  between  the  sexes,  as  in  the  size  of  the  canine  teeth,  in 
the  development  and  color  of  the  hair,  and  especially  in  the  color 
of  the  naked  parts  of  the  skin,  than  in  mankind. 

All  the  secondary  sexual  characters  of  man  are  highly  variable, 
even  within  the  limits  of  the  same  race;  and  they  differ  much  in 
the  several  races.  These  two  rules  hold  good  generally  through- 
out the  animal  kingdom.  In  the  excellent  observations  made  on 
board  the  Novara,^-  the  male  Australians  were  found  to  exceed 
the  females  by  only  65  millim.  in  height,  whilst  with  the  Javans 
the  average  excess  was  218  millim.;  so  that  in  this  latter  race  the 
difference  in  height  between  the  sexes  is  more  than  thrice  as  great 
as  with  the  Australians.  Numerous  measurements  were  carefully 
made  of  the  stature,  the  circumference  of  the  neck  and  chest,  the 

10  Mr.  Blyth  informs  me  that  he  has  only  seen  one  instance  of  the 
beard,  whiskers,  &c.,  in  a  monkey  becoming-  white  with  old  age,  as 
is  so  commonly  the  case  with  us.  This,  however,  occurred  in  an  aged 
Macacus  cynomolgus,  kept  in  confinement,  whose  moustaches  were 
"remarkably  long  and  human-like."  Altogether  this  old  monkey  pre- 
sented a  ludicrous  resemblance  to  one  of  the  reigning  monarchs  of 
Europe,  after  whom  he  was  universally  nick-named.  In  certain  races 
of  man  the  hair  on  the  head  hardly  ever  becomes  gray;  thus  Mr.  D. 
Forbes-  has  never,  as  he  informs  me,  seen  an  instance  with  the 
Aymaras  and  Quichuas  of  S.  America. 

^  This  is  the  case  with  the  females  of  several  species  of  Hylobates, 
see  GeofErey  St.-Hilaire  and  F.  Cuvier,  'Hist.  Nat.  des  Mamm.'  tom.  i. 
See,  also,  on  H.  lar.  'Penny  Cyclopedia,'  vol.  ii.  pp.  149,  150. 

12  The  results  were  deduced  by  Dr.  Weisbach  from  the  measure- 
ments made  by  Drs.  K.  Scherzer  and  Schwarz,  see  'Reise  der  Novara: 
Anthropolog.  Theil,'  1867,  ss.  216,  231,  234,  236^  239,  269. 


MAN— SEXUAL  DIFFERENCES.  555 

length  of  the  back-bone  and  of  the  arms,  in  various  races;  and 
nearly  all  these  measurements  show  that  the  males  differ  much 
more  from  one  another  than  do  the  females.  This  fact  indicates 
that,  as  far  as  these  characters  are  concerned,  it  is  the  male  which 
has  been  chiefly  modified,  since  the  several  races  diverged  from 
their  common  stock. 

The  development  of  the  beard  and  the  hairiness  of  the  body  dif- 
fer remarkably  in  the  men  of  distinct  races,  and  even  in  different 
tribes  or  families  of  the  same  race.  We  Europeans  see  this 
amongst  ourselves.  In  the  Island  of  St.  Kilda,  according  to  Mar- 
tin," the  men  do  not  acquire  beards  until  the  age  of  thirty  or  up- 
wards, and  even  then  the  beards  are  very  thin.  On  the  Europseo- 
Asiatic  continent,  beards  prevail  until  we  pass  beyond  India; 
though  with  the  natives  of  Ceylon  they  are  often  absent,  as  was 
noticed  in  ancient  times  by  Diodorus."  Eastward  of  India  beards 
disappear,  as  with  the  Siamese,  Malays,  Kalmucks,  Chinese,  and 
Japanese;  nevertheless  the  Ainos,'^  who  inhabit  the  northernmost 
islands  of  the  Japan  Archipelago,  are  the  hairiest  men  in  the 
world.  With  negroes  the  beard  is  scanty  or  wanting,  and  they 
rarely  have  whiskers;  in  both  sexes  the  body  is  frequently  almost 
destitute  of  fine  down.^*^  On  the  other  hand,  the  Papuans  of  the 
Malay  Archipelago,  who  are  nearly  as  black  as  negroes,  possess 
well-developed  beards."  In  the  Pacific  Ocean  the  inhabitants  of 
the  Fiji  Archipelago  have  large  bushy  beards,  whilst  those  of  the 
not  distant  archipelagoes  of  Tonga  and  Samoa  are  beardless;  but 
these  men  belong  to  distinct  races.  In  the  Ellice  group  all  the  in- 
habitants belong  to  the  same  race;  yet  on  one  island  alone,  namely 
Nunemaya,  "the  men  have  splendid  beards;"  whilst  on  the  other 
islands  "they  have,  as  a  rule,  a  dozen  straggling  hairs  for  a 
"beard."i« 

Throughout  the  great  American  continent  the  men  may  be 
said  to  be  beardless;  but  in  almost  all  the  tribes  a  few  short  hairs 
are  apt  to  appear  on  the  face,  especially  in  old  age.  With  the 
tribes  of  North  America,  Catlin  estimates  that  eighteen  out  of 

13  'Voyage  to  St.  Kilda'  (3rd  edit.  1753),  p.  37. 
1*  Sir  J.  E.  Tennent,   'Ceylon,'  vol.  ii.  1859,  p.  107. 

■^5  Quatrefages,  'Revue  des  Cours  Scientifiques,'  Aug.  29,  1868,  p.  630; 
Vogt,  'Lectures  on  Man,'  Eng.  translat.  p.  127. 

16  On  the  beards  of  negroes,  Vogt,  'Lectures,'  &c.  p.  127;  Waitz, 
'Introduct.  to  Anthropology,'  Engl,  translat.  1863,  vol.  i.  p.  96.  It  is 
remarkable  that  in  the  United  States  ('Investigations  in  Military  and 
Anthropological  Statistics  of  American  Soldiers,'  1869,  p.  569)  the  pure 
negroes  and  their  crossed  offspring  seem  to  have  bodies  almost  as 
hairy  as  Europeans. 

17  Wallace,   'The  Malay  Arch.'  vol.  ii.  1869,  p.  178. 

18  Dr.  J.  Barnard  Davis  On  Oceanic  Races,  in  'Anthropolog.  Review,' 
April,  1870,  pp.  185,  191. 


556  THE  DESCENT  OF  MAN, 

twenty  men  are  completely  destitute  by  nature  of  a  beard;  but 
occasionally  there  may  be  seen  a  man,  who  has  neglected  to  plucK 
out  the  hairs  at  puberty,  with  a  soft  beard  an  inch  or  two  in 
length.  The  Guaranys  of  Paraguay  differ  from  all  the  surround- 
ing tribes  in  having  a  small  beard,  and  even  some  hair  on  the 
body,  but  no  whiskers."  I  am  informed  by  Mr.  D.  Forbes,  who 
particularly  attended  to  this  point,  that  the  Aymaras  and  Quichuas 
of  the  Cordillera  are  remarkably  hairless,  yet  in  old  age  a  few 
straggling  hairs  occasionally  appear  on  the  chin.  The  men  of 
these  two  tribes  have  very  little  hair  on  the  various  parts  of  tiie 
body  where  hair  grows  abundantly  in  Europeans,  and  the  women 
have  none  on  the  corresponding  parts.  The  hair  on  the  hea-l, 
however,  attains  an  extraordinary  length  in  both  sexes,  often 
reaching  almost  to  the  ground;  and  this  is  likewise  the  case  with 
some  of  the  N.  American  tribes.  In  the  amount  of  hair,  and  in  the 
general  shape  of  the  body,  the  sexes  of  the  American  aborigines  do 
not  differ  so  much  from  each  other,  as  in  most  other  races.-*'  This 
fact  is  analogous  with  what  occurs  with  some  closely  allied  mon- 
keys; thus  the  sexes  of  the  chimpanzee  are  not  as  different  as 
those  of  the  orang  or  gorilla.-^ 

In  the  previous  chapters  we  have  seen  that  with  mammals, 
birds,  fishes,  insects,  &c.,  many  characters,  which  there  is  every 
reason  to  believe  were  primarily  gained  through  sexual  selection 
by  one  sex,  have  been  transferred  to  the  other.  As  this  same  form 
of  transmission  has  apparently  prevailed  much  with  mankind, 
it  will  save  useless  repetition  if  we  discuss  the  origin  of  characters 
peculiar  to  the  male  sex  together  with  certain  other  characters 
common  to  both  sexes. 

Law  of  Battle.  — With  savages,  for  instance  the  Australians,  the 
women  are  the  constant  cause  of  war  both  between  members  of 
the  same  tribe  and  between  distinct  tribes.  So  no  doubt  it  was 
in  ancient  times;  "nam  fuit  ante  Helenam  mulier  teterrima  belli 
"causa."  With  some  of  the  North  American  Indians,  the  contest 
is  reduced  to  a  system.   That  excellent  observer,  Hearne,^^  says:  — 

i»  Catlin,  'North  American  Indians,'  3rd  edit.  1842,  vol.  ii.  p.  227.  On 
the  Guaranys,  see  Azara,  'Voyages  dans  rAmerique  Merid.'  torn.  ii. 
1809,  p.  58;    also,  Reng-ger,  'Saugethiere  von  Paraguay,'  s.  3. 

20  Prof,  and  Mrs.  Agassiz  ('Journey  in  Brazil,'  p.  530)  remark  that  the 
sexes  of  the  American  Indians  differ  less  than  those  of  the  negroes 
and  of  the  higher  races.  See,  also,  Rengger,  ibid.  p.  3,  on  the 
Guaranys. 

21  Rutimeyer,  'Die  Grenzen  der  Thierwelt;  eine  Betrachtung  zu  Dar- 
win's Lehre,'  18G8,  s.  54. 

22  'A  Journey  from  Prince  of  Wales  Fort,'  8vo.  edit.  Dublin,  1796, 
p.  104.  Sir  J.  Lubbock  ('Origin  of  Civilization,'  1870,  p.  69)  gives  other 
and  similar  cases  in  North  America.  For  the  Guanas  of  S.  America, 
see  Azara,  'Voyages,'  &c.,  torn.  ii.  p.  94. 


MAN— LAW  OF  BATTLE.  557 

"It  has  ever  been  the  custom  among  these  people  for  the  men  to 
"wrestle  for  any  woman  to  whom  they  are  attached;  and,  of 
"course,  the  strongest  party  always  carries  off  the  prize.  A  weali 
"man,  unless  he  be  a  good  hunter,  and  well-beloved,  is  seldom  per- 
"mitted  to  keep  a  wife  that  a  stronger  man  thinks  worth  his  no- 
"tice.  This  custom  prevails  throughout  all  the  tribes,  and  causes 
"a  great  spirit  of  emulation  among  their  youth,  who  are  upon  all 
"occasions,  from  their  childhood,  trying  their  strength  and  skill 
"in  wrestling."  With  the  Guanas  of  South  America,  Azara  states 
that  the  men  rarely  marry  till  twenty  years  old  or  more,  as  before 
that  age  they  cannot  conquer  their  rivals. 

Other  similar  facts  could  be  given;  bilt  even  if  we  had  no  evi- 
dence on  this  head,  we  might  feel  almost  sure,  from  the  analogy 
of  the  higher  Quadrumana,-^  that  the  law  of  battle  had  prevailed 
with  man  during  the  early  stages  of  his  development.  The  occa- 
sional appearance  at  the  present  day  of  canine  teeth  which  project 
above  the  others,  with  traces  of  a  diastema  or  open  space  for  the 
reception  of  the  opposite  canines,  is  in  all  probability  a  case  of  re- 
version to  a  former  state,  when  the  progenitors  of  man  were  pro- 
vided with  these  weapons,  like  so  many  existing  male  Quadru- 
mana.  It  was  remarked  in  a  former  chapter  that  as  man  gradually 
became  erect,  and  continually  used  his  hands  and  arms  for  fighting 
with  sticks  and  stones,  as  well  as  for  the  other  purposes  of  life,  he 
would  have  used  his  jaws  and  teeth  less  and  less.  The  Jaws  to- 
gether with  their  muscles,  would  then  have  been  reduced  through 
disuse,  as  would  the  teeth  through  the  not  well  understood  princi- 
ples of  correlation  and  economy  of  growth;  for  we  everywhere  see 
that  parts,  which  are  no  longer  of  service,  are  reduced  in  size.  By 
such  steps  the  original  inequality  between  the  jaws  and  teeth  in 
the  two  sexes  of  mankind  would  ultimately  have  been  obliterated. 
The  case  is  almost  parallel  with  that  of  many  male  Ruminants,  in 
which  the  canine  teeth  have  been  reduced  to  mere  rudiments,  or 
have  disappeared,  apparently  in  consequence  of  the  development  of 
horns.  As  the  prodigious  difference  between  the  skulls  of  the  two 
sexes  in  the  orang  and  gorilla  stands  in  close  relation  with  the 
development  of  the  immense  canine  teeth  in  the  males,  we  may 
infer  that  the  reduction  of  the  jaws  and  teeth  in  the  early  male 
progenitors  of  man  must  have  led  to  a  most  striking  and  favorable 
change  in  his  appearance. 

There  can  be  little  doubt  that  the  greater  size  and  strength  of 
man,  in  comparison  with  woman,  together  with  his  broa'der  shoul- 
ders, more  developed  muscles,  rugged  outline  of  body,  his  greater 
courage  and  pugnacity,  are  all  due  in  chief  part  to  inheritance 

-2  On  the  fighting-  of  the  male  gorillas,  see  Dr.  Savage,  in  'Boston 
Journal  of  Nat.  Hist.'  vol.  v.  1847,  p.  423.  On  Presbytia  entellus,  see 
the  'Indian  Field,'  1859,  p.  146. 


558  THE  DESCENT  OF  MAN. 

from  his  half-human  male  ancestors.  These  characters  would, 
however,  have  been  preserved  or  even  augmented  during  the  long 
ages  of  man's  savagery,  by  the  success  of  the  strongest  and  boldest 
men,  both  in  the  general  struggle  for  life  and  in  their  contests 
for  wives;  a  success  which  would  have  ensured  their  leaving  a 
more  numerous  progeny  than  their  less  favored  brethren.  It  is 
not  probable  that  the  greater  strength  of  man  was  primarily  ac- 
quired through  the  inherited  e:ffiects  of  his  having  worked  harder 
than  woman  for  his  own  subsistence  and  that  of  his  family;  for  the 
women  in  all  barbarous  nations  are  compelled  to  work  at  least 
as  hard  as  the  men.  With  civilized  people  the  arbitrament  of 
battle  for  the  possession  of  the  women  has  long  ceased;  on  the 
other  hand,  the  men,  as  a  general  rule,  have  to  work  harder  than 
the  women  for  their  joint  subsistence,  and  thus  their  greater 
strength  will  have  been  kept  up. 

Difference  in  the  Mental  Powers  of  the  two  Sexes.  — ^With  respect 
to  differences  of  this  nature  between  man  and  woman,  it  is  prob- 
able that  sexual  selection  has  played  a  highly  important  part.  1 
am  aware  that  some  writers  doubt  whether  there  is  any  such  in- 
herent difference;  but  this  is  at  least  probable  from  the  analogy  of 
the  lower  animals  which  present  other  secondary  sexual  charac- 
ters. No  one  disputes  that  the  bull  differs  in  disposition  from  the 
cow,  the  wild-boar  from  the  sow,  the  stallion  from  the  mare,  and, 
as  is  well  known  to  the  keepers  of  menageries,  the  males  of  the 
larger  apes  from  the  females.  Woman  seems  to  differ  from  man 
in  mental  disposition,  chiefly  in  her  greater  tenderness  and  less 
selfishness;  and  this  holds  good  even  with  savages,  as  shown  by  a 
well-known  passage  in  Mungo  Park's  Travels,  and  by  statements 
made  by  many  other  travelers.  Woman,  owing  to  her  maternal 
instincts,  displays  these  qualities  towards  her  infants  in  an  emi- 
nent degree;  therefore  it  is  likely  that  she  would  often  extend 
them  towards  her  fellow-creatures.  Man  is  the  rival  of  other  men; 
he  delights  in  competition,  and  this  leads  to  ambition  which  passes 
too  easily  into  selfishness.  These  latter  qualities  seem  to  be  his 
natural  and  unfortunate  birthright.  It  is  generally  admitted  that 
with  woman  the  powers  of  intuition,  of  rapid  perception,  and  per- 
haps of  imitation,  are  more  strongly  marked  than  in  man;  but 
some,  at  least,  of  these  faculties  are  characteristic  of  the  lower 
races,  and  therefore  of  a  past  and  lower  state  of  civilization. 
T  The  chief  distinction  in  the  intellectul  powers  of  the  two  sexes 
V'  is  shown  by  man's  attaining  to  a  higher  eminence,  in  whatever  he 
takes  up,  than  can  woman — ^whether  requiring  deep  thought,  rea- 
son, or  imagination,  or  merely  the  use  of  the  senses  and  hands. 
If  two  lists  were  made  of  the  most  eminent  men  and  women  in 
poetry,  painting,  sculpture,  music  (inclusive  both  of  composition 
and  performance),  history,  science,  and  philosophy,  with  half- a- 


MENTAL   POWERS  OF  MAN  AND  WOMAN.  559 

dozen  names  under  each  subject,  the  two  lists  would  not  bear  com- 
parison. "We  may  also  infer,  from  the  law  of  the  deviation  from 
averages,  so  well  illustrated  by  Mr.  Galton,  in  his  work  on  'Hered- 
itary genius,'  that  if  men  are  capable  of  a  decided  pre-eminence 
over  women  in  many  subjects,  the  average  of  mental  power  in 
man  must  be  above  that  of  woman. 

Amongst  the  half-human  progenitors  of  man,  and  amongst  sav- 
ages, there  have  been  struggles  between  the  males  during  many 
generations  for  the  possession  of  the  females.  But  mere  bodily 
strength  and  size  would  do  little  for  victory,  unless  associated 
with  courage,  perseverance,  and  determined  energy.  With  social 
animals,  the  young  males  have  to  pass  through  many  a  contest  be- 
fore they  win  a  female,  and  the  older  males  have  to  retain  their  fe- 
males by  renewed  battles.  They  have,  also,  in  the  case  of  man- 
kind, to  defend  their  females,  as  well  as  their  young,  from  enemies 
of  all  kinds,  and  to  hunt  for  their  joint  subsistence.  But  to  avoid 
enemies  or  to  attack  them  with  success,  to  capture  wild  animals, 
and  to  fashion  weapons,  requires  the  aid  of  the  higher  mental 
faculties,  namely,  observation,  reason,  invention,  or  imagination. 
These  various  faculties  will  thus  have  been  continually  put  to  the 
test  and  selected  during  manhood;  they  will,  moreover,  have  been 
strengthened  by  use  during  this  same  period  of  life.  Conse- 
quently, in  accordance  with  the  principle  often  alluded  to,  we 
might  expect  that  they  Y^^ould  at  least  tend  to  be  transmitted 
chiefly  to  the  male  offspring  at  the  corresponding  period  of  man- 
hood. 

Now,  when  two  men  are  put  into  competition,  or  a  man  with  a 
woman,  both  possessed  of  every  mental  quality  in  equal  perfec- 
tion, save  that  one  has  higher  energy,  perseverance,  and  courage, 
the  latter  will  generally  become  more  eminent  in  every  pursuit, 
and  will  gain  the  ascendancy.^*  He  may  be  said  to  possess 
genius — for  genius  has  been  declared  by  a  great  authority  to  be 
patience;  and  patience,  in  this  sense,  means  unflinching,  un- 
daunted perseverance.  But  this  view  of  genius  is  perhaps  de- 
ficient; for  without  the  higher  powers  of  the  imagination  and 
reason,  no  eminent  success  can  be  gained  in  many  subjects.  These 
latter  faculties,  as  well  as  the  former,  will  have  been  developed  in 
man,  partly  through  sexual  selection, — that  is,  through  the  contest 
of  rival  males,  and  partly  through  natural  selection, — that  is, 
from  success  in  the  general  struggle  for  life;  and  as  in  both  cases 
the  struggle  will  have  been  during  maturity,  the  characters  gained 
will  have  been  transmitted  more  fully  to  the  male  than  to  the  fe- 


2*  J.  Stuart  Mill  remarks  ('The  Subjection  of  Women,'  1869,  p.  122), 
"The  things  in  which  man  most  excels  woman  are  those  which  require 
"most  plodding,  and  long  hammering  at  single  thoughts."  What  is 
this  but  energy  and  perseverance? 


560  THE  DESCENT  OF  MAN. 

male  offspring.  It  accords  in  a  striking  manner  with  this  view 
of  the  modification  and  re-inforcement  of  many  of  our  mental 
faculties  by  sexual  selection,  that,  firstly,  they  notoriously  under- 
go a  considerable  change  at  puberty ,2=  and,  secondly,  that  eunuchs 
remain  throughout  life  inferior  in  these  same  qualities.  Thus 
man  has  ultimately  become  superior  to  woman.  It  is,  indeed, 
fortunate  that  the  law  of  the  equal  transmission  of  characters  to 
both  sexes  prevails  with  mammals;  otherwise  it  is  probable  that 
man  would  have  become  as  superior  in  mental  endowment  to 
woman,  as  the  peacock  is  in  ornamental  plumage  to  the  peahen. 

It  must  be  borne  in  mind  that  the  tendency  in  characters  ac- 
quired by  either  sex  late  in  life,  to  be  transmitted  to  the  same  sex 
at  the  same  age,  and  of  early  acquired  characters  to  be  transmitted 
to  both  sexes,  are  rules  which,  though  general,  do  not  always  hold. 
If  they  always  held  good,  we  might  conclude  (but  I  here  exceed  my 
proper  bounds)  that  the  inherited  effects  of  the  early  education  of 
boys  and  girls  would  be  transmitted  equally  to  both  sexes;  so  that 
the  present  inequality  in  mental  power  between  the  sexes  would 
not  be  effaced  by  a  similar  course  of  early  training;  nor  can  it 
have  been  caused  by  their  dissimilar  early  training.  In  order  that 
woman  should  reach  the  same  standard  as  man,  she  ought,  when 
nearly  adult,  to  be  trained  to  energy  and  perseverance,  and  to  have 
her  reason  and  imagination  exercised  to  the  highest  point;  and 
then  she  would  probably  transmit  these  qualities  chiefly  to  her 
adult  daughters.  Ail  women,  however,  could  not  be  thus  raised, 
unless  during  many  generations  those  who  excelled  in  the  above 
robust  virtues  were  married,  and  produced  offspring  in  larger 
numbers  than  other  women.  As  before  remarked  of  bodily 
strength,  although  men  do  not  now  fight  for  their  wives,  and  this 
form  of  selection  has  passed  away,  yet  during  manhood,  they  gen- 
erally undergo  a  severe  struggle  in  order  to  maintain  themselves 
and  their  families;  and  this  will  tend  to  keep  up  or  even  increase 
their  mental  powers,  and,  as  a  consequence,  the  present  inequality 
between  the  sexes.^^ 

Voice  and  Musical  Powers.  — In  some  species  of  Quadrumana 
there  is  a  great  difference  between  the  adult  sexes,  in  the  power 
of  their  voices  and  in  the  development  of  the  vocal  organs;  and 

25  Maudsley,  'Mind  and  Body,'  p.  31. 

23  An  observation  by  Vogt  bears  on  this  subject:  he  says,  "It  is  a 
"remarkable  circumstance,  that  the  difference  between  the  sexes,  as 
"reg-ards  the  cranial  cavity,  increases  with  the  development  of  the 
"race,  so  that  the  male  European  excels  much  more  the  female,  than 
"the  negro  the  negress.  Welcker  confirms  this  statement  of  Huschke 
"from  his  measurements  of  negro  and  German  skulls."  But  Vogt  ad- 
mits ('Lectures  on  Man,'  Eng.  translat.  1864,  p.  81)  that  more  observa- 
tions are  requisite  on  this  point. 


MAN— MUSICAL  POWERS.  561 

man  appears  to  have  inherited  this  difference  from  his  early  pro- 
genitors. His  vocal  cords  are  about  one-third  longer  than  in  v/o- 
man,  or  than  in  hoys;  and  emasculation  produces  the  same  effect 
on  him  as  on  the  lower  animals,  for  it  "arrests  that  prominent 
"growth  of  the  thyroid,  &c.,  which  accompanies  the  elongation  or 
"the  cords."-'^  With  respect  to  the  cause  of  this  difference  between 
the  sexes,  I  have  nothing  to  add  to  the  remarks  in  the  last  chapter 
on  the  probable  effects  of  the  long-continued  use  of  the  vocal  or- 
gans by  the  male  under  the  excitement  of  love,  rage  and  jealousy. 
According  to  Sir  Duncan  Gibb,"''  the  voice  and  the  form  of  the 
larynx  differ  in  the  different  races  of  mankind;  but  with  the  Tar- 
tars, Chinese,  &c.,  the  voice  of  the  male  is  said  not  to  differ  so 
much  from  that  of  the  female,  as  in  most  other  races. 

The  capacity  and  love  for  singing  or  music,  though  not  a  sexual 
character  in  man,  must  not  here  be  passed  over.  Although  the 
sounds  emitted  by  animals  of  all  kinds  serve  many  purposes,  a 
strong  case  can  be  made  out,  that  the  vocal  organs  were  primarily 
used  and  perfected  in  relation  to  the  propagation  of  the  species. 
Insects  and  some  few  spiders  are  the  lowest  animals  which  volun- 
tarily produce  any  sound;  and  this  is  generally  effected  by  the 
aid  of  beautifully  constructed  stridulating  organs,  which  are  often 
confined  to  the  males.  The  sounds  thus  produced  consist,  I  believe 
in  all  cases,  of  the  same  note,  repeated  rhythmically; ^^  and  this  is 
sometimes  pleasing  even  to  the  ears  of  man.  The  chief  and,  in 
some  cases,  exclusive  purpose  appears  to  be  either  to  call  or 
charm  the  opposite  sex. 

The  sounds  produced  by  fishes  are  said  in  some  cases  to  be  made 
only  by  the  males  during  the  breeding-season.  All  the  air-breath- 
ing Vertebrata  necessarily  possess  an  apparatus  for  inhaling  and 
expelling  air,  with  a  pipe  capable  of  being  closed  at  one  end. 
Hence  when  the  primeval  members  of  this  class  were  strongly  ex- 
cited and  their  muscles  violently  contracted,  purposeless  sounds 
would  almost  certainly  have  been  produced;  and  these,  if  they 
proved  in  any  way  serviceable,  might  readily  have  been  modified 
or  intensified  by  the  preservation  of  properly  adapted  variations. 
The  lowest  Vertebrates  which  breathe  air  are  Amphibians;  and  of 
these,  frogs  and  toads  possess  vocal  organs,  which  are  incessantly 
used  during  the  breeding-season,  and  which  are  often  more  highly 
developed  in  the  male  than  in  the  female.  The  male  alone  of  the 
tortoise  utters  a  noise,  and  this  only  during  the  season  of  love. 
Male  alligators  roar  or  bellow  during  the  same  season.  Every  one 
knows  how  much  birds  use  their  vocal  organs  as  a  means  of  court - 


27  Owen,  'Anatomy  of  Vertebrates,'  vol.  lii.  p.  603. 
23  'Journal  of  the  Anthropolog-.  Soc'  April,  1869,  p.  Ivii.  and  Ixvi. 
29  Dr.  Scudder,  'Notes  on  Stridulation,'  in  'Proc.  Boston  Soc.  of  Nat. 
Hist.'  vol.  xl.  April,  1868. 
3T 


562  THE  DESCENT  OF  MAN. 

ship;  and  some  species  likewise  perform  what  may  be  called  in- 
strumental music. 

In  the  class  of  Mammals,  with  which  we  are  here  more  par- 
ticularly concerned,  the  males  of  alraost  all  the  species  use  their 
voices  during  the  breeding-season  much  more  than  at  any  other 
time;  and  some  are  absolutely  mute  excepting  at  this  season.  With 
other  species  both  sexes,  or  only  the  females,  use  their  voices  as  a 
love-call.  Considering  these  facts,  and  that  the  vocal  organs  of 
some  quadrupeds  are  much  more  largely  developed  in  the  male 
than  in  the  female,  either  permanently  or  temporarily  during  the 
breeding-season;  and  considering  that  in  most  of  the  lower  classes 
the  sounds  produced  by  the  males,  serve  not  only  to  call  but  to  ex- 
cite or  allure  the  female,  it  is  a  surprising  fact  that  we  have  not 
as  yet  any  good  evidence  that  these  organs  are  used  by  male  mam- 
mals to  charm  the  females.  The  American  Mycetes  caraya  per- 
haps forms  an  exception,  as  does  the  Hylobates  agilis,  an  ape  al- 
lied to  man.  This  gibbon  has  an  extremely  loud  but  musical 
voice.  Mr.  Waterhouse  states.^"  "It  appeared  to  me  that  in  as- 
"cending  and  descending  the  scale,  the  intervals  were  always  ex- 
"actly  half-tones;  and  I  am  sure  that  the  highest  note  was  the 
"exact  octave  to  the  lowest.  The  quality  of  the  notes  is  very 
"musical;  and  I  do  not  doubt  that  a  good  violinist  would  be  a  bio 
*'to  give  a  correct  idea  of  the  gibbon's  composition,  excepting  as 
"regards  its  loudness."  Mr.  Waterhouse  then  gives  the  notes. 
Professor  Owen,  who  is  a  musicidin,  confirms  the  foregoing  state- 
ment, and  remarks,  though  erroneously,  that  this  gibbon  "alone 
"of  brute  mammals  may  be  said  to  sing."  It  appears  to  be  much 
excited  after  its  performance.  Unfortunately,  its  habits  have 
never  been  closely  observed  in  a  state  of  nature;  but  from  the 
analogy  of  other  animals,  it  is  probable  that  it  uses  its  musical 
powers  more  especially  during  the  season  of  courtship. 

This  gibbon  is  not  the  only  species  in  the  genus  which  sings,  for 
my  son,  Francis  Darwin,  attentively  listened  in  the  Zoological  Gar- 
dens to  H.  ieuciscus  whilst  singing  a  cadence  of  three  notes,  in 
true  musical  intervals  and  with  a  clear  musical  tone.  It  is  a  more 
surprising  fact  that  certain  rodents  utter  musical  sounds.  Sing- 
ing mice  have  often  been  mentioned  and  exhibited,  biit  imposture 
has  commonly  been  suspected.  We  have,  however,  at  last  a  clear 
account  by  a  well-known  observer,  the  Rev.  S.  Lockwood,  of  the 
musical  powers  of  an  American  species,  the  Hesperomys  cognatus, 
belonging  to  a  genus  distinct  from  that  of  the  English  mouse. 
This  little  animal  was  kept  in  confinement,  and  the  performance 

30  Given  in  W.  C,  L.  Martin's  'General  Introduct.  to  Nat.  Hist,  ol 
Mamm.  Animals,'  1841,  p.  432;  Owen,  'Anatomy  of  Vertebrates,'  vol.  iii. 
p.  600. 

31  The  'American  Naturalist,'  1871,  p.  761- 


MAN— MUSICAL  POWERS.  563 

was  repeatedly  heard.  In  one  of  the  two  chief  songs,  "the  last 
"bar  would  frequently  be  prolonged  to  two  or  three;  and  she 
"would  sometimes  change  from  C  sharp  and  D,  to  C  natural  and 
"D,  then  warble  on  these  two  notes  awhile,  and  wind  up  with  a 
"quick  chirp  on  C  sharp  and  D.  The  distinctness  between  the 
"semitones  was  very  marked,  and  easily  appreciable  to  a  good 
"ear."  Mr.  Lockv/ood  gives  both  songs  in  musical  notation;  and 
?,dds  that  though  this  little  mouse  "had  no  ear  for  time,  yet  she 
"would  keep  to  the  key  of  B  (two  flats)  and  strictly  in  a  major 

"key." "Her  soft  clear  voice  falls  an  octave  with  all  the  pre- 

"cision  possible;  then  at  the  wind  up,  it  rises  again  into  a  very 
"quick  trill  on  C  sharp  and  D." 

A  critic  has  asked  how  the  ears  of  man,  and  he  ought  to  have 
added  of  other  animals,  could  have  been  adapted  by  selection  so  as 
to  distinguish  musical  notes.  But  this  question  shows  some  confu- 
sion on  the  subject;  a  noise  is  the  sensation  resulting  from  the 
co-existence  of  several  aerial  "simple  vibrations"  of  various 
periods,  each  of  which  intermits  so  frequently  that  its  separate 
existence  cannot  be  perceived.  It  is  only  in  the  want  of  continuity 
of  such  vibrations,  and  in  their  want  of  harmony  inter  se,  that 
a  noise  differs  from  a  musical  note.  Thus  an  ear  to  be  capable 
of  discriminating  noises— and  the  high  importance  of  this  power 
to  all  animals  is  admitted  by  every  one — must  be  sensitive  to  musi- 
cal notes.  We  have  evidence  of  this  capacity  even  low  dov/n  in 
the  animal  scale:  thus  Crustaceans  are  provided  with  auditory 
hairs  of  different  lengths,  which  have  been  seen  to  vibrate  when 
the  proper  musical  notes  are  struck.^-  As  stated  in  a  previous 
chapter,  similar  observations  have  been  made  on  the  hairs  of  the 
antennas  of  gnats.  It  has  been  positively  asserted  by  good  obser- 
vers that  spiders  are  attracted  by  music.  It  is  also  well  known 
that  some  dogs  howl  when  hearing  particular  tones.^  Seals  ap- 
parently appreciate  music,  and  their  fondness  for  it  "was  well 
"known  to  the  ancients,  and  is  often  taken  advantage  of  by  the 
"hunters  at  the  present  day."^* 

Therefore,  as  far  as  the  mere  perception  of  musical  notes  is  con- 
cerned, there  seems  no  special  difficulty  in  the  case  of  man  or  of 
any  other  animal.  Helmholtz  has  explained  on  physiological  prin- 
ciples why  concords  are  agreeable,  and  discords  disagreeable  to  the 
human  ear;  but  we  are  little  concerned  with  these,  as  music  in  har- 


32  Helmholtz,  'Theorie  Phys,  de  la  Musique,'  1868,  p.  187. 

33  Several  accounts  have  been  published  to  this  effect.  Mr,  Peach 
writes  to  me  that  he  has  repeatedly  found  that  an  old  dog-  of  his 
howls  when  B  flat  is  sounded  on  the  flute,  and  to  no  other  note.  I 
may  add  another  instance  of  a  dog  always  whining-,  when  one  note 
on  a  concertina,  which  was  oiit  of  tune,  was  played. 

s^  Mr.  R.  Brown,  in  'Proc.  Zool.  Soc'  1868,  p.  410. 


564  THE  DESCENT  OF  MAN. 

inony  is  a  late  invention.  We  are  more  concerned  with  melody, 
and  here  again,  according  to  Helmholtz,  it  is  intelligible  why  the. 
notes  of  our  musical  scale  are  used.  The  ear  analyzes  all  sounds 
into  their  component  "simple  vibrations,"  although  we  are  not 
conscious  of  this  analysis.  In  a  musical  note  the  lowest  in  pitch 
of  these  is  generally  predominant,  and  the  others  which  are  less 
marked  are  the  octave,  the  twelfth,  the  second  octave,  &c.,  all 
harmonies  of  the  fundamental  predominant  note;  any  two  notes 
of  our  scale  have  many  of  these  harmonic  over-tones  in  common. 
It  seems  pretty  clear  then,  that  if  an  animal  always  wished  to  sing 
precisely  the  same  song,  he  would  guide  himself  by  sounding  those 
notes  in  succession,  which  possess  many  over- tones  in  common — 
that  is,  he  would  choose  for  his  song,  notes  which  belong  to  our 
musical  scale. 

But  if  it  be  further  asked  why  musical  tones  in  a  certain  order 
and  rhythm  give  man  and  other  animals  pleasure,  we  can  no 
more  give  the  reason  than  for  the  pleasantness  of  certain  tastes 
and  smells.  That  they  do  give  pleasure  of  some  kind  to  animals, 
we  may  infer  from  their  being  produced  during  the  season  of 
courtship  by  many  insects,  spiders,  fishes,  amphibians,  and  birds; 
for  unless  the  females  w^ere  able  to  appreciate  such  sounds  and 
were  excited  or  charmed  by  them,  the  persevering  efforts  of  the 
males,  and  the  complex  structures  often  possessed  by  them  alone, 
would  be  useless;  and  this  it  is  impossible  to  believe. 

Human  song  is  generally  admitted  to  be  the  basis  or  origin  of 
instrumental  music.  As  neither  the  enjoyment  nor  the  capacity 
of  producing  musical  notes  are  faculties  of  the  least  use  to  man 
in  reference  to  his  daily  habits  of  life,  they  must  be  ranked 
amongst  the  most  mysterious  with  which  he  is  endowed.  They 
are  present,  though  in  a  very  rude  condition,  in  men  of  all  races, 
even  the  most  savage;  but  so  different  is  the  taste  of  the  several 
races,  that  our  music  gives  no  pleasure  to  savages,  and  their  music 
is  to  us  in  most  cases  hideous  and  unmeaning.  Dr.  Seemann,  in 
some  interesting  remarks  on  this  subject,^^  "doubts  whether  even 
"amongst  the  nations  of  Western  Europe,  intimately  connected 
"as  they  are  by  close  and  frequent  intercourse,  the  music  of  the 
"one  is  interpreted  in  the  same  sense  by  the  others.  By  traveling 
"eastwards  we  find  that  there  is  certainly  a  different  language  of 
"music.  Songs  of  joy  and  dance-accompaniments  are  no  longer, 
"as  with  us,  in  the  major  keys,  but  always  in  the  minor."  Whether 
or  not  the  half-human  progenitors  of  man  possessed,  like  the  sing- 
ing gibbons,  the  capacity  of  producing,  and  therefore  no  doubt  of 

35  'Journal  of  Anthropolog.  Soc'  Oct.  1870,  p.  civ.  See,  also,  the 
several  later  chapters  in  Sir  John  Lubbock's  'Prehistoric  Times,"  sec- 
ond edition,  1869,  which  contain  an  admirable  account  of  the  habits  of 
savages. 


MAN— MUSICAL  POWERS.  S65 

appreciating  musical  notes,  we  know  that  man  possessed  these 
faculties  at  a  very  remote  period.  M.  Lartet  has  described  two 
flutes,  made  out  of  the  bones  and  horns  of  the  reindeer,  found  in 
caves  together  with  flint  tools  and  the  remains  of  extinct  animals. 
The  arts  of  singing  and  of  dancing  are  also  very  ancient,  and  are 
now  practiced  by  all  or  nearly  all  the  lowest  races  of  man.  Poetry, 
which  may  be  considered  as  the  offspring  of  song,  is  likewise  so 
ancient,  that  many  persons  have  felt  astonished  that  it  should 
have  arisen  during  the  earliest  ages  of  which  we  have  any  record. 
We  see  that  the  musical  faculties,  which  are  not  wholly  deficient 
in  any  race,  are  capable  of  prompt  and  high  development,  for  Hot- 
tentots and  Negroes  have  become  excellent  musicians,  although  in 
their  native  countries  they  rarely  practice  anything  that  we 
should  consider  music.  Schweinfurth,  however,  was  pleased  with 
some  of  the  simple  melodies  which  he  heard  in  the  interior  of 
Africa.  But  there  is  nothing  anomalous  in  the  musical  faculties 
lying  dormant  in  man:  some  species  of  birds  vv^hich  never  natur- 
ally sing,  can  without  much  difficulty  be  taught  to  do  so;  thus  a 
house-sparrow  has  learnt  the  song  of  a  linnet.  As  these  two 
species  are  closely  allied,  and  belong  to  the  order  of  Insessores, 
which  includes  nearly  all  the  singing-birds  in  the  world,  it  is  pos- 
sible that  a  progenitor  of  the  sparrow  may  have  been  a  songster. 
It  is  more  remarkable  that  parrots,  belonging  to  a  group  distinct 
from  the  Insessores,  and  having  differently  constructed  vocal  or- 
gans, can  be  taught  not  only  to  speak,  but  to  pipe  or  whistle  tunes 
invented  by  man,  so  that  they  must  have  some  musical  capacity. 
Nevertheless  it  would  be  very  rash  to  assume  that  parrots  are 
descended  from  some  ancient  form  which  was  a  songster.  Many 
cases  could  be  advanced  of  organs  and  instincts  originally  adapted 
for  one  purpose,  having  been  utilized  for  some  distinct  purpose.^*" 
Hence  the  capacity  for  high  musical  development,  which  the  sav- 
age races  of  man  possess,  may  be  due  either  to  the  practice  by  our 
semi-human  progenitors  of  some  rude  form  of  music,  or  simply  to 
their  having  acquired  the  proper  vocal  organs  for  a  different  pur- 
pose. But  in  this  latter  case  we  must  assume,  as  in  the  above  in- 
stance of  parrots,  and  as  seems  to  occur  with  many  animals,  that 
they  already  possessed  some  sense  of  melody. 

30  Since  this  chapter  was  printed,  I  have  seen  a  valuable  article  by- 
Mr.  Chauncey  Wright  ('North  Amer.  Review,'  Oct.  1870,  page  293), 
who,  in  discussing  the  above  subject,  remarks,  "There  are  many  con- 
"sequences  of  the  ultimate  laws  or  uniformities  of  nature,  through 
"which  the  acquisition  of  one  useful  power  will  bring  with  it  many 
"resulting  advantages  as  well  as  limiting  disadvantages,  actual  or  pos- 
"sible,  which  the  principle  of  utihty  may  not  have  comprehended  in 
■■■  tts  action."  As  I  have  attempted  to  show  in  an  early  chapter  of  this 
work,  this  principle  has  an  important  bearing  on  the  acQvcisitiou  by 
man  of  some  of  his  mental  characteristics. 

37 


566  THE  DESCENT  OF  MAN. 

Music  arouses  in  us  various  emotions,  but  not  the  more  terrible 
ones  of  horror,  fear,  rage,  &c.  It  awakens  the  gentler  feelings  ot 
tenderness  and  love,  which  readily  pass  into  devotion.  In  the 
Chinese  annals  it  is  said,  "Music  hath  the  power  of  making  heaven 
"descend  upon  earth."  It  likewise  stirs  up  in  us  the  sense  of 
triumph  and  the  glorious  ardor  for  war.  These  powerful  and  min- 
gled feelings  may  well  give  rise  to  the  sense  of  sublimity.  We  can 
concentrate,  as  Dr.  Seemann  observes,  greater  intensity  of  feel- 
ing in  a  single  musical  note  than  in  pages  of  writing.  It  is  prob- 
able that  nearly  the  same  emotions,  but  much  weaker  and  far  less 
complex,  are  felt  by  birds  when  the  male  pours  forth  his  full 
volume  of  song,  in  rivalry  with  other  males,  to  captivate  the  fe- 
male. Love  is  still  the  commonest  theme  of  our  songs.  As  Her- 
bert Spencer  remarks,  "music  arouses  dormant  sentiments  of 
"which  we  had  not  conceived  the  possibility,  and  do  not  know  the 
"meaning;  or,  as  Richter  says,  tells  us  of  things  we  have  not  seen 
"and  shall  not  see."  Conversely,  when  vivid  emotions  are  felt 
and  expressed  by  the  orator,  or  even  in  common  speech,  musical 
cadences  and  rhythm  are  instinctively  used.  The  negro  in  Africa 
when  excited  often  bursts  forth  in  song;  "another  will  reply  in 
"song,  while  the  company,  as  if  touched  by  a  musical  wave,  mur- 
"mur  a  chorus  in  perfect  unison."-''  Even  monkeys  express  strong 
feelings  in  different  tones — anger  and  impatience  by  low, — fear 
and  pain  by  high  notes.^^  The  sensations  and  ideas  thus  excited 
in  us  by  music,  or  expressed  by  the  cadences  of  oratory,  appear 
from  their  vagueness,  yet  depth,  like  mental  reversions  to  the 
emotions  and  thoughts  of  a  long-past  age. 

All  these  facts  with  respect  to  music  and  impassioned  speech 
become  intelligible  to  a  certain  extent,  if  we  may  assume  that 
musical  tones  and  rhythm  were  used  by  our  half-human  ancestors 
during  the  season  of  courtship,  when  animals  of  all  kinds  are  ex- 
cited not  only  by  love,  but  by  the  strong  passions  of  jealousy, 
rivalry,  and  triumph.  From  the  deeply-laid  principle  of  inherited 
associations,  musical  tones  in  this  case  would  be  likely  to  call  up 
vaguely  and  indefinitely  the  strong  emotions  of  a  long-past  age. 
As  we  have  every  reason  to  suppose  that  articulate  speech  is  one 
of  the  latest,  as  it  certainly  is  the  highest,  of  the  arts  acquired  by 
man,  and  as  the  instinctive  power  of  producing  musical  notes  and 
rhythms  is  developed  low  down  in  the  animal  series,  it  would  be 
altogether  opposed  to  the  principle  of  evolution,  if  we  were  to  ad- 
mit that  man's  musical  capacity  has  been  developed  from  the  tones 
used  in  impassioned  speech.  We  must  suppose  that  the  rhythms 
and  cadences  of  oratory  are  derived  from  previously  developed 

s''  Winwood  Reade,  'The  Martyrdom  of  Man,'  1872,  p.  441,  and  'African 
Sketch  Book,'  1873,  vol.  ii.  p.  313. 
^  Reng-ger,  'Saugethiere  von  Paraguay,'  s.  49. 


MAN— MUSICAL  POWERS.  567 

musical  powers.^^  "We  can  tlius  understand  how  it  is  that  music, 
dancing,  song,  and  poetry  are  such  very  ancient  arts.  We  may 
go  even  further  than  this,  and,  as  remarked  in  a  former  chapter, 
believe  that  musical  sounds  afforded  one  of  the  bases  for  the  de- 
velopment of  the  language.*" 

As  the  males  of  several  quadrumanous  animals  have  their  vocal 
organs  much  more  developed  than  in  the  females,  and  as  a  gibbon, 
one  of  the  anthropomorphous  apes,  pours  forth  a  whole  octave  of 
musical  notes  and  may  be  said  to  sing,  it  appears  probable  that 
the  progenitors  of  man,  either  the  males  or  females  or  both  sexes, 
before  acquiring  the  power  of  expressing  their  mutual  love  in  ar- 
ticulate language,  endeavored  to  charm  each  other  with  musical 
notes  and  rhythm.  So  little  is  known  about  the  use  of  the  voice 
by  the  Quadrumana  during  the  season  of  love,  that  we  have  no 
means  of  judging  whether  the  habit  of  singing  was  first  acquired 
by  our  male  or  female  ancestors.  Women  are  generally  thought 
to  possess  sweeter  voices  than  men,  and  as  far  as  this  serves  as 
any  guide,  we  may  infer  that  they  first  acquired  musical  powers 
in  order  to  attract  the  other  sex.*^  But  if  so,  this  must  have  oc- 
curred long  ago,  before  our  ancestors  had  become  sufiaciently  hu- 
man to  treat  and  value  their  women  merely  as  useful  slaves.  The 
impassioned  orator,  bard,  or  musician,  when  with  his  varied  tones 
and  cadences  he  excites  the  strongest  emotions  in  his  hearers,  lit- 
tle suspects  that  he  uses  the  same  means  by  which  his  half-human 


3»  See  the  very  interesting-  discussion  on  the  'Origin  and  Function  of 
Music,'  by  Mr.  Herbert  Spencer,  in  his  collected  'Essays,'  1858,  p.  359. 
Mr.  Spencer  comes  to  an  exactly  opposite  conclusion  to  that  at  which 
I  have  arrived.  He  concludes,  as  did  Diderot  formerly,  that  the  ca- 
dences used  in  emotional  speech  afford  the  foundation  from  which 
music  has  been  developed;  whilst  I  conclude  that  musical  notes  and 
rhythm  were  first  acquired  by  the  male  or  female  prog-enitors  of  man- 
kind for  the  sake  of  charming-  the  opposite  sex.  Thus  musical  tones 
became  firmly  associated  with  some  of  the  strongest  passions  an  ani- 
mal is  capable  of  feeling,  and  are  consequently  used  instinctively,  or 
through  association,  when  strong  emotions  are  expressed  in  speech. 
Mr.  Spencer  does  not  offer  any  satisfactory  explanation,  nor  can  I, 
why  high  or  deep  notes  should  be  expressive,  both  with  man  and  the 
lower  animals,  of  certain  emotions.  Mr.  Spencer  gives  also  an  in- 
teresting discussion  on  the  relations  between  poetry,  recitative,  and 
song. 

*o  I  find  in  Lord  Monboddo's  'Origin  of  Language,'  vol.  i.  (1774),  p. 
469,  that  Dr.  Blacklock  likewise  thought  "that  the  first  language 
"among  men  was  music,  and  that  before  our  ideas  were  expressed  by 
"articulate  sounds,  they  were  communicated  by  tones,  varied  ac- 
"cording  to  different  degrees  of  gravity  and  acuteness." 

"■  See  an  interesting  discussion  on  this  subject  by  Hackel,  'Gene^ 
relle  Morph,'  B.  ii.  1866,  s.  246. 


568  THE  DESCENT  OF  MAN. 

ancestors  long  ago  aroused  each  other's  ardent  passions,  during 
their  courtship  and  rivalry. 

The  Influence  of  Beauty  in  determining  the  Marriages  of  Man- 
Mnd. — In  civilized  life  man  is  largelj^  but  by  no  means  exclu- 
sively, influenced  in  the  choice  of  his  wife  by  external  appear- 
ance; but  we  are  chiefly  concerned  with  primeval  times,  and  our 
only  means  of  forming  a  judgment  on  this  subject  is  to  study 
the  habits  of  existing  semi-civilized  and  savage  nations.  If  it  can 
be  shown  that  the  men  of  different  races  prefer  women  having 
various  characteristics,  or  conversely  with  the  women,  we  have 
then  to  inquire  whether  such  choice,  continued  during  many  gen- 
erations, would  produce  any  sensible  effect  on  the  race,  either  on 
one  sex  or  both  according  to  the  form  of  inheritance  w^hich  has 
prevailed. 

It  will  be  well  first  to  show  in  some  detail  that  savages  pay  the 
greatest  attention  to  their  personal  appearance.--  That  they  have 
a  passion  for  ornament  is  notorious;  and  an  English  philosopher 
goes  so  far  as  to  maintain,  that  clothes  were  first  made  for  orna- 
ment and  not  for  v/armth.  As  Professor  Waitz  remarks,  "how- 
"ever  poor  and  miserable  man  is,  he  finds  a  pleasure  in  adorning 
"himself."  The  extravagance  of  the  naked  Indians  of  South 
America  in  decorating  themselves  is  shown  "by  a  man  of  large 
"stature  gaining  with  difficulty  enough  by  the  labor  of  a  fortnight 
"to  procure  in  exchange  the  chica  necessary  to  paint  himself 
"red."''^  The  ancient  barbarians  of  Europe  during  the  Reindeer 
period  brought  to  their  caves  any  brilliant  or  singular  objects 
which  they  happened  to  find.  Savages  at  the  present  day  every- 
where deck  themselves  v/ith  plumes,  necklaces,  armlets,  ear-rings, 
&c.  They  paint  themselves  in  the  most  diversified  manner.  "If 
"painted  nations,"  as  Humboldt  observes,  "had  been  examined 
"with  the  same  attention  as  clothed  nations,  it  would  have  been 
"perceived  that  the  most  fertile  imagination  and  the  most  muta- 

*2  A  full  and  excellent  account  of  the  manner  in  which  savages  in  all 
parts  of  the  world  ornament  themselves,  is  given  by  the  Italian  trav- 
eler, Prof.  Mantegazza,  'Rio  de  la  Plata,  Viaggi  e  Studi,'  1S87,  pp.  525- 
545:  all  the  following  statements,  when  other  references  are  not 
given,  are  taken  from  this  work.  See,  also,  "Waitz,  'Introduct.  to  An- 
thropolog.'  Eng.  transl.  vol.  i.  1863,  p.  275,  et  passim.  Lawrence  also 
gives  very  full  details  in  his  'Lectures  on  Physiology,'  1822.  Since  this 
chapter  was  written  Sir  J.  Lubbock  has  published  his  'Origin  of 
Civilization,'  1870,  in  which  there  is  an  interesting  chapter  on  the 
present  subject,  and  from  which  (pp.  42,  48)  I  have  taken  some  facts 
about  savages  dyeing  their  teeth  and  hair,  and  piercing  their  teeth. 

^  Humboldt,  'Personal  Narrative,'  Eng.  Translat.  vol.  iv.  p.  515;  on 
the  imagination  shown  in  painting  the  body,  p.  522;  on  modifying  the 
form  of  the  calf  of  the  leg,  p.  466. 


MAN— LOVE    OF    ORNAMENT.  569 

"ble  caprice  have  created  the  fashions  of  painting,  as  well  as 
"those  of  garments." 

In  one  part  of  Africa  the  eyelids  are  colored  black;  in  another 
the  nails  are  colored  yellow  or  purple.  In  many  places  the  hair 
is  dyed  of  various  tints.  In  different  countries  the  teeth  are 
stained  black,  red,  blue,  &c.,  and  in  the  Malay  Archipelago  it  is 
thought  shameful  to  have  white  teeth  "like  those  of  a  dog."  Not 
one  great  country  can  be  named,  from  the  Polar  regions  in  the 
north  to  New  Zealand  in  the  south,  in  which  the  aborigines  do 
not  tattoo  themselves.  This  practice  was  follov/ed  by  the  Jews 
of  old,  and  by  the  ancient  Britons.  In  Africa  some  of  the  na- 
tives tattoo  themselves,  but  it  is  a  much  more  common  prac- 
tice to  raise  protuberances  by  rubbing  salt  into  incisions  made 
in  various  parts  of  the  body;  and  these  are  considered  by  the 
inhabitants  of  Kordofan  and  Darfur  "to  be  great  personal  at- 
"tractions."  In  the  Arab  countries  no  beauty  can  be  perfect 
until  the  cheeks  "or  temples  have  been  gashed."^*  In  South 
America,  as  Humboldt  remarks,  "a  mother  would  be  accused  of 
"culpable  indifference  "towards  her  children,  if  she  did  not  em- 
"ploy  artificial  means  to  shape  the  calf  of  the  leg  after  the  fashion 
"of  the  country."  In  the  Old  and  New  Worlds  the  shape  of  the 
skull  was  formerly  modified  during  infancy  in  the  most  extraor- 
dinary manner,  as  is  still  the  case  in  many  places,  and  such  de- 
formities are  considered  ornamental.  For  instance,  the  savages 
of  Colombia^  deem  a  much  flattened  head  "an  essential  point 
"of  beauty." 

The  hair  is  treated  with  especial  care  in  various  countries;  it 
is  allowed  to  grow  to  full  length,  so  as  to  reach  to  the  ground,  or 
is  combed  into  "a  compact  frizzled  mop,  which  is  the  Papuan's 
"pride  and  glory."^^  In  Northern  Africa  "a  man  requires  a  period 
"of  from  eight  to  ten  years  to  perfect  his  coiffure."  With  other 
nations  the  head  is  shaved,  and  in  parts  of  South  America  and 
Africa  even  the  eyebrows  and  eyelashes  are  eradicated.  The  na- 
tives of  the  Upper  Nile  knock  out  the  four  front  teeth,  saying 
that  they  do  not  wish  to  resemble  brutes.  Further  south,  the 
Batokas  knock  out  only  the  two  upper  incisors,  which,  as  Liv- 
ingstone^^ remarks,  gives  the  face  a  hideous  appearance,  owing 
to  the  prominence  of  the  lower  jaw;    but  these  people  think  the 


*4  'The  Nile  Tributaries,'  1867;  'The  Albert  N'yanza,'  1866,  vol.  i. 
p.  218. 

45  Quoted  by  Prichard,  'Phys.  Hist,  of  Mankind,'  4th  edit.  vol.  i.  1851, 
p.  321. 

46  On  the  Papuans,  Wallace,  'The  Malay  Archipelag-o,'  vol.  ii.  p.  445, 
On  the  coiffure  of  the  Africans,  Sir  S.  Baker,  'The  Albert  N'yanza,' 
vol.  i.  p.  210. 

4'='  'Travels,'  p.  533. 


570  THE  DESCENT  OF  MAN. 

presence  of  the  incisors  most  unsigMly,  and  on  beholding  some 
Europeans,  cried  out,  "Look  at  the  great  teeth!"  The  chief 
Sebituani  tried  in  vain  to  alter  this  fashion.  In  various  parts  of 
Africa  and  in  the  Malay  Archipelago  the  natives  file  the  incisors 
into  points  like  those  of  a  saw,  or  pierce  them  with  holes,  into 
which  they  insert  studs. 

As  the  face  with  us  is  chiefly  admired  for  its  beauty,  so  with 
savages  it  is  the  chief  seat  of  mutilation.  In  all  quarters  of  the 
world  the  septum  and  more  rarely  the  wings  of  the  nose  are 
pierced;  rings,  sticks,  feathers,  and  other  ornaments  being  in- 
serted into  the  holes.  The  ears  are  everywhere  pierced  and  sim- 
ilarly ornamented,  and  with  the  Botocudos  and  Lenguas  of  South 
America  the  hole  is  gradually  so  much  enlarged  that  the  lower 
edge  touches  the  shoulder.  In  North  and  South  America  and  in 
Africa  either  the  upper  or  lower  lip  is  pierced;  and  with  the 
Botocudos  the  hole  in  the  lower  lip  is  so  large  that  a  disc  of  wood, 
four  inches  in  diameter,  is  placed  in  it.  Mantegazza  gives  a  curious 
account  of  the  shame  felt  by  a  South  American  native  and  of 
the  ridicule  which  he  excited,  when  he  sold  his  tembeta,— the 
large  colored  piece  of  wood  which  is  passed  through  the  hole.  In 
Central  Africa  the  women  perforate  the  lower  lip  and  wear  a  crys- 
tal, which,  from  the  movement  of  the  tongue,  has  "a  wriggling 
"motion,  indescribably  ludicrous  during  conversation."  The  wife 
of  the  chief  of  Latooka  told  Sir  S.  Baker*^  that  Lady  Baker  "would 
"be  much  improved  if  she  would  extract  her  four  front  teeth 
"from  the  lower  jaw,  and  wear  the  long  pointed  polished  crystal 
"in  her  under  lip."  Further  south  with  the  Makalolo,  the  upper 
lip  is  perforated,  and  a  large  metal  and  bamboo  ring,  called  a 
pelele,  is  worn  in  the  hole.  "This  caused  the  lip  in  one  case 
"to  project  two  inches  beyond  the  tip  of  the  nose;  and  when 
"the  lady  smiled  the  contraction  of  the  muscles  elevated  it  over 
"the  eyes.  'Why  do  the  women  wear  these  things?'  the  ven- 
"erable  chief,  Chinsurdi,  was  asked.  Evidently  surprised  at  such 
"a  stupid  question,  he  replied,  'For  beauty!  They  are  the  only 
"  'beautiful  things  women  have;  men  have  beards,  women  have 
"  'none.  What  kind  of  a  person  would  she  be  without  the  pelele? 
"  'She  would  not  be  a  woman  at  all  with  a  mouth  like  a  man, 
"  'but  no  beard.'  "*" 

Hardly  any  part  of  the  body,  which  can  be  unnaturally  modi- 
fied, has  escaped.  The  amount  of  suffering  thus  caused  must 
have  been  extreme,  for  many  of  the  operations  require  several 
years  for  their  completion,  so  that  the  idea  of  their  necessity 
must  be  imperative.    The  motives  are  various;  the  men  paint  their 

48  'The  Albert  N'yanza,'  1866,  vol.  i.  p.  217. 

49  Livingstone,     'British     Association,'     1860;      report    given    in    the 
♦Athenaeum.'  -Tuly  7,  1860.  p.  29. 


MAN— BEAUTY.  571 

bodies  to  make  themselves  appear  terrible  in  battle;  certain  muti- 
lations are  connected  with  religious  rites,  or  they  mark  the  age 
of  puberty,  or  the  rank  of  the  man,  or  they  serve  to  distinguish 
the  tribes.  Amongst  savages  the  same  fashions  prevail  for  long 
periods,'^"  and  thus  mutilations,  from  whatever  cause  first  made, 
soon  come  to  be  valued  as  distinctive  marks.  But  self-adornment, 
vanity,  and  the  admiration  of  others,  seem  to  be  the  commonest 
motives.  In  regard  to  tattooing,  I  was  told  by  the  missionaries 
in  New  Zealand,  that  when  they  tried  to  persuade  some  girls  to 
give  up  the  practice,  they  answered,  "We  must  just  have  a  few 
"lines  on  our  lips;  else  when  we  grow  old  we  shall  be  so  very 
"ugly."  With  the  men  of  New  Zealand,  a  most  capable  judge^^  says, 
"to  have  fine  tattooed  faces  was  the  great  ambition  of  the  young, 
"both  to  render  themselves  attractive  to  the  ladies,  and  conspicu- 
"ous  in  war."  A  star  tattooed  on  the  forehead  and  a  spot  on  the 
chin  are  thought  by  the  women  in  one  part  of  Africa  to  be 
irresistible  attractions.^^  In  most,  but  not  all  parts  of  the  world, 
the  men  are  more  ornamented  than  the  women,  and  often  in  a 
different  manner;  sometimes,  though  rarely,  the  women  are 
hardly  at  all  ornamented.  As  the  women  are  made  by  savages  to 
perform  the  greatest  share  of  the  work,  and  as  they  are  not  al- 
lowed to  eat  the  best  kinds  of  food,  so  it  accords  with  the  char- 
acteristic selfishness  of  man  that  they  should  not  be  allowed  to 
obtain,  or  use  the  finest  ornaments.  Lastly,  it  is  a  remarkable 
fact,  as  proved  by  the  foregoing  quotations,  that  the  same  fash- 
ions in  modifying  the  shape  of  the  head,  in  ornamenting  the  hair, 
in  painting,  tattooing,  in  perforating  the  nose,  lips,  or  ears,  in 
removing  or  filing  the  teeth,  &c.,  now  prevail,  and  have  long 
prevailed,  in  the  most  distant  quarters  of  the  world.  It  is  ex- 
tremely improbable  that  these  practices,  followed  by  so  many 
distinct  nations,  should  be  due  to  tradition  from  any  common 
source.  They  indicate  the  close  similarity  of  the  mind  of  man, 
to  whatever  race  he  may  belong,  just  as  do  the  almost  universal 
habits  of  dancing,  masquerading,  and  making  rude  pictures. 

Having  made  these  preliminary  remarks  on  the  admiration 
felt  by  savages  for  various  ornaments,  and  for  deformities  most 
unsightly  in  our  eyes,  let  us  see  how  far  the  men  are  attracted 
by  the  appearance  of  their  women,  and  what  are  their  ideas  of 
beauty.     I  have  heard  it  maintained  that  savages  are  quite  in- 

E^o  Sir  S.  Baker  (ibid.  vol.  i.  p.  210)  speaking-  of  the  natives  of  Central 
Africa  says,  "every  tribe  has  a  distinct  and  unchanging-  fashion  for 
"dressing  the  hair."  See  Agassiz  ('Journey  in  Brazil,'  1868,  p.  318)  on 
the  invariability  of  the  tattooing-  of  the  Amazonian  Indians. 

51  Rev.  R.  Taylor,  'New  Zealand  and  its  Inhabitants,'  1855,  p.  152. 

^2  Manteg-azza,  'Viaggi  e  Studi,'  p.  542. 


572  THE  DESCENT  OP  MAN. 

different  about  the  beauty  of  their  women,  valuing  them  solely 
as  slaves;  it  may  therefore  be  well  to  observe  that  this  conclusion 
does  not  at  all  agree  with  the  care  which  the  women  take  in 
ornamenting  themselves,  or  with  their  vanity.  Burcheir'^  gives 
an  amusing  account  of  a  Bush-woman  who  used  as  much  grease, 
red  ochre,  and  shining  powder  "as  would  have  ruined  any  but  a 
"very  rich  husband."  She  displayed  also  "much  vanity  and  too 
"evident  a  consciousness  of  her  superiority."  Mr.  Winwood  Reade 
informs  me  that  the  negroes  of  the  West  Coast  often  discuss  the 
beauty  of  their  women.  Some  competent  observers  have  attrib- 
uted the  fearfully  common  practice  of  infanticide  partly  to  the 
desire  felt  by  the  women  to  retain  their  good  loolis.*^*  In  several 
regions  the  women  wear  charms  and  use  love-philters  to  gain 
the  affections  of  the  men;  and  Mr.  Brown  enumerates  four  plants 
used  for  this  purpose  by  the  women  of  North-Western  America.^^ 

Hearne,'''°  an  excellent  observer,  who  lived  many  years  with  the 
American  Indians,  says,  in  speaking  of  the  women,  "Ask  a  North- 
"ern  Indian  what  is  beauty,  and  he  will  answer,  a  broad  flat  face, 
"small  eyes,  high  cheek-bones,  three  or  four  broad  black  lines 
"across  e-ach  cheek,  a  low  forehead,  a  large  broad  chin,  a  clumsy 
"hook  nose,  a  tav/ny  hide,  and  breasts  hanging  dov/n  to  the  belt." 
Pallas,  who  visited  the  northern  parts  of  the  Chinese  empire, 
nays  "those  women  are  preferred  who  have  the  Mandschu  form; 
'"that  is  to  say,  a  broad  face,  high  cheek-bones,  very  broad  noses, 
".',nd  enormous  ears;"^^  and  Vogt  remarks  that  the  obliquity  of 
the  eye,  which  is  proper  to  the  Chinese  and  Japanese,  is  exag- 
gerated in  their  pictures  for  the  purpose,  as  it  "seems,  of  exhibit- 
"ing  its  beauty,  as  contrasted  with  the  eye  of  the  red-haired  bar- 
"barians."  It  is  well  known,  as  Hue  repeatedly  remarks,  that 
the  Chinese  of  the  interior  think  Europeans  hideous,  with  their 
white  skins  and  prominent  noses.  The  nose  is  far  from  being 
too  prominent,  according  to  our  ideas,  in  the  natives  of  Ceylon; 
yet  "the  Chinese  in  the  seventh  century,  accustomed  to  the  flat 
"features  of  the  Mongol  races,  were  surprised  at  the  prominent 
"noses  of  the  Cingalese;  and  Thsang  described  them  as  having 
"  'the  beak  of  a  bird,  with  the  body  of  a  man.'  " 

Finlayson,  after  minutely  describing  the  people  of  Cochin  China, 

53  'Travels  in  S.  Africa,'  1824,  vol.  i.  p.  414. 

^*  See,  for  references,  Gerland  'Ueber  das  Aussterben  der  Naturvol- 
ker,'  1868,  s.  51,  53,  55;    also  Azara  'Voyages,'  «&;c.  torn.  ii.  p.  116. 

55  On  the  vegetable  productions  used  by  the  North-Western  Ameri- 
can Indians,   'Pliarmaceutical  Journal,'  vol.  x. 

^«  'A  Journey  from  Prince  of  Wales  Fort,'  8vo.  edit.  1796,  p.  89. 

^7  Quoted  by  Prichard,  'Phys.  Hist,  of  Mankind,'  3rd  edit.  vol.  iv. 
1844,  p.  519;  Vog-t,  'Lectures  on  Man,'  Eng-.  translat.  p.  129.  On  the 
opinion  of  the  Chinese  on  the  Cingalese,  E.  Tennent,  'Ceylon,'  1859, 
vol.  ii.  p.  107. 


MAN— BEAtll^Tr.  g7^ 

says  that  their  rounded  heads  and  faces  are  the  chief  charac- 
teristics; and,  he  adds,  "the  roundness  of  the  whole  countenance 
"is  more  striking  in  the  women,  who  are  reckoned  beautiful  in 
"proportion  as  they  display  this  form  of  face."  The  Siamese 
have  small  noses  with  divergent  nostrils,  a  wide  mouth,  rather 
thick  lips,  a  remarkably  large  face,  with  very  high  and  broad 
cheek-bones.  It  is,  therefore,  not  wonderful  that  "beauty,  accord- 
"ing  to  our  notion  is  a  stranger  to  them.  Yet  they  consider  their 
"own  females  to  be  much  more  beautiful  than  those  of  Europe. "^^ 

It  is  well  known  that  with  many  Hottentot  women  the  posterior 
part  of  the  body  projects  in  a  wonderful  manner;  they  are  steatop- 
ygous;  and  Sir  Andrew  Smith  is  'certain  that  this  peculiarity  is 
greatly  admired  by  the  men.'^^  He  once  saw  a  woman  who  was 
considered  a  beauty,  and  she  was  so  immensely  developed  behind, 
that  when  seated  on  level  ground  she  could  not  rise,  and  had 
to  push  herself  along  until  she  came  to  a  slope.  Some  of  the 
women  in  various  negro  tribes  have  the  same  peculiarity;  and, 
according  to  Burton,  the  Somal  men  "are  said  to  choose  their 
"wives  by  ranging  them  in  a  line,  and  by  picking  her  out  who 
"projects  farthest  a  tergo.  Nothing  can  be  more  hateful  to  a 
"negro  than  the  opposite  form.'"'" 

With  respect  to  color,  the  negroes  rallied  Mungo  Park  on  the 
whiteness  of  his  skin  and  the  prominence  of  his  nose,  both  of 
which  they  considered  as  "unsightly  and  unnatural  conforma- 
"tions."  He  in  return  praised  the  glossy  jet  of  their  skins  and  the 
lovely  depression  of  their  noses;  this  they  said  was,  "honey- 
"mouth,"  nevertheless  they  gave  him  food.  The  African  Moors, 
also,  "knitted  their  brows  and  seemed  to  shudder"  at  the  white- 
ness of  his  skin.  On  the  eastern  coast,  the  negro  boys  when  they 
saw  Burton,  cried  out  "Look  at  the  white  man;  does  he  not  look 
"like  a  white  ape?"  On  the  western  coast,  as  Mr.  Winwood 
Reade  informs  me,  the  negroes  admire  a  very  black  skin  more 
than  one  of  a  lighter  tint.  But  their  horror  of  whiteness  may  be 
attributed,  according  to  this  same  traveler,  partly  to  the  belief 
held  by  most  negroes  that  demons  and  spirits  are  white,  and 
partly  to  their  thinking  it  a  sign  of  ill-health. 

The  Banyai  of  the  more  southern  part  of  the  continent  are 
negroes,  but  "a  great  many  of  them  are  of  a  light  coffee-and-milk 

Bsprichard,  as  taken  from  Crawfurd  and  Finlayson,  'Phys.  Hist,  of 
Mankind,'  vol.  iv.  pp.  534,  535. 

59  Idem  illustrissimus  viator  dixit  mihi  praecinctorium  vel  tabulam 
foeminae,  quod  nobis  teterrimum  est,  quondam  permagno  aestimari  ab 
hominibus  in  hac  gente.  Nunc  res  mutata  est,  et  censent  talem  con- 
forniationem  minime  optandam  esse. 

^  'The  Anthropological  Reviev^,'  November,  1864,  p.  237.  For  addi- 
tional references,  see  Waitz,  'Introduct.  to  Anthropology,'  Eng.  trans- 
lat.  1863,  vol.  1.  p.  105. 


574  THE  DESCENT  OF  MAN. 

"color,  and,  indeed,  this  color  is  considered  handsome  through- 
"out  the  whole  country;"  so  that  here  we  have  a  different  stand- 
ard of  taste.  With  the  Kafirs,  who  differ  much  from  negroes, 
"the  skin,  except  among  the  tribes  near  Delagoa  Bay,  is  not  usually 
"black,  the  prevailing  color  being  a  mixture  of  black  and  red, 
"the  most  common  shade  being  chocolate.  Dark  complexions,  as 
"being  most  common  are  naturally  held  in  the  highest  esteem. 
"To  be  told  that  he  is  light-colored,  or  like  a  white  man,  would 
"be  deemed  a  very  poor  compliment  by  a  Kafir.  I  have  heard 
"of  one  unfortunate  man  who  was  so  very  fair  that  no  girl  would 
"marry  him."  One  of  the  titles  of  the  Zulu  king  is  "You  who  are 
"black."^^  Mr.  Galton,  in  speaking  to  me  about  the  natives  of 
S.  Africa,  remarked  that  their  ideas  of  beauty  seem  very  different 
from  ours;  for  in  one  tribe  two  slim,  slight,  and  pretty  girls  were 
not  admired  by  the  natives. 

Turning  to  other  quarters  of  the  world;  in  Java,  a  yellow,  not 
a  white  girl,  is  considered,  according  to  Madame  Pf eiffer,  a  beauty. 
A  man  of  Cochin  China  "spoke  with  contempt  of  the  wife  of  the 
"English  Ambassador,  that  she  had  white  teeth  like  a  dog,  and  a 
"rosy  color  like  that  of  potato-flowers."  We  have  seen  that  the 
Chinese  dislike  our  white  skin,  and  that  the  N,  Americans  admire 
"a  tawny  hide."  In  S.  America,  the  Yuracaras,  who  inhabit  the 
wooded,  damp  slopes  of  the  eastern  Cordillera,  are  remarkably 
pale-colored,  as  their  name  in  their  own  language  expresses; 
nevertheless  they  consider  European  women  as  very  inferior  to 
their  own.®^ 

In  several  of  the  tribes  of  North  America  the  hair  on  the  head 
grows  to  a  wonderful  length;  and  Catlin  gives  a  curious  proof 
how  much  this  is  esteemed,  for  the  chief  of  the  Crows  was 
elected  to  this  office  from  having  the  longest  hair  of  any  man  in 
the  tribe,  namely  ten  feet  and  seven  inches.  The  Aymaras  and 
Quichuas  of  S.  America,  likewise  have  very  long  hair;  and  this, 
as  Mr.  D.  Forbes  informs  me,  is  so  much  valued  as  a  beauty, 
that  cutting  it  off  was  the  severest  punishment  which  he  could 
inflict  on  them.  In  both  the  Northern  and  Southern  halves  of 
the  continent  the  natives  sometimes  increase  the  apparent  length 
of  their  hair  by  weaving  into  it  fibrous  substances.  Although 
the  hair  on  the  head  is  thus  cherished,  that  on  the  face  is  con- 

eiMungo  Park's  'Travels  in  Africa.'  4to.  1816,  pp.  53,  131.  Burton's 
statement  is  quoted  by  Schaaffhausen,  'Archiv.  fur  Anthropolog.'  1866, 
s.  163.  On  the  Banyai,  Livingstone,  'Travels,'  p.  64.  On  the  Kaflra, 
the  Rev.  J.  Shooter,  'The  Kafirs  of  Natal  and  the  Zulu  Country,'  1857, 

p.  1. 

62  por  the  Javans  and  Cochin-Chinese,  see  Waitz,  'Introduct.  to 
Anthropology",'  Eng.  translat.  vol.  i.  p.  305.  On  the  Tura-caras,  A. 
d'Orligny,  as  quoted  in  Prichard,  'Phys.  Hist,  of  Mankind,'  vol.  v.  3rd 
eOit.  p.  476. 


MAN— BEAUTY.  575 

sidered  by  the  North  American  Indians  *'as  very  vulgar,"  and 
every  hair  is  carefully  eradicated.  This  practice  prevails  through- 
out the  American  continent  from  Vancouver's  Island  in  the  north 
to  Tierra  del  Fuego  in  the  south.  When  York  Minster,  a  Fuegian 
on  hoard  the  'Beagle,'  was  taken  hack  to  his  country,  the  natives 
told  him  he  ought  to  pull  out  the  few  short  hairs  on  his  face.  They 
also  threatened  a  young  missionary,  who  was  left  for  a  time  with 
them,  to  strip  him  naked,  and  pluck  the  hairs  from  his  face  and 
body,  yet  he  was  far  from  being  a  hairy  man.  This  fashion  is 
carried  so  far  that  the  Indians  of  Paraguay  eradicate  their 
eyebrows  and  eyelashes,  saying  that  ithey  do  not  wish  to  be  like 
horses.^^ 

It  is  remarkable  that  throughout  the  world  the  races  which 
are  almost  completely  destitute  of  a  beard,  dislike  hairs  on  the 
face  and  body,  and  take  pains  to  eradicate  them.  The  Kalmucks 
are  beardless,  and  they  are  well  known,  like  the  Americans,  to 
pluck  out  all  straggling  hairs;  and  so  it  is  with  the  Polynesians, 
some  of  the  Malays,  and  the  Siamese.  Mr.  Veitch  states  that  the 
Japanese  ladies  "all  objected  to  our  whiskers,  considering  them 
"very  ugly,  and  told  us  to  cut  them  off,  and  be  like  Japanese 
"men."  The  New  Zealanders  have  short,  curled  beards;  yet 
they  formerly  plucked  out  the  hairs  on  the  face.  They  had  a 
saying  that  "there  is  no  woman  for  a  hairy  man;"  but  it  would 
appear  that  the  fashion  has  changed  in  New  Zealand,  perhaps 
owning  to  the  presence  of  Europeans,  and  I  am  assured  that 
beards  are  now  admired  by  the  Maories.^ 

On  the  other  hand,  bearded  races  admire  and  greatly  value 
their  beards;  among  the  Anglo-Saxons  every  part  of  the  body 
had  a  recognized  value;  "the  loss  of  the  beard  being  estimated 
"at  twenty  shillings,  while  the  breaking  of  a  thigh  was  fixed  at 
"only  twelve."*^^  In  the  East  men  swear  solemnly  by  their  beards. 
We  have  seen  that  Chinsurdi,  the  chief  of  the  Makalolo  in  Africa, 
thought  that  beards  were  a  great  ornament.  In  the  Pacific  the 
Fijian's  beard  is  "profuse  and  bushy,  and  is  his  greatest  pride;" 
whilst  the  inhabitants  of  the  adjacent  archipelagoes  of  Tonga  and 
Samoa  are  "beardless,  and  abhor  a  rough  chin."     In  one  island 

63  'North  American  Indians,'  by  G.  Catlin,  3rd  edit.  1842,  vol.  i.  p. 
49;  vol.  ii.  p.  227.  On  the  natives  of  Vancouver's  Island,  see  Sproat, 
•Scenes  and  Studies  of  Savage  Life,'  1868,  p.  25.  On  the  Indians  of 
Paraguay,  Azara,  'Voyages,'  torn.  ii.  p.  105. 

^  On  the  Siamese,  Prichard,  ibid.  vol.  iv.  p.  533.  On  the  Japanese, 
Veitch  in  'Gardeners'  Chronicle,'  1860,  p.  1104.  On  the  New  Zealanders, 
Mantegazza,  'Viaggi  e  Studi,'  1867,  p.  526.  For  the  other  nations  men- 
tioned, see  references  in  Lawrence,  'Lectures  on  Physiology,'  &c.  1822, 
p.  272. 

65  Lubbock,  'Origin  of  Civilization,'  1870,  p.  32L 


576  THE   DESCENT  OF  MAN. 

alone  of  the  Ellice  group  "the  men  are  heavily  bearded,  and  not 
"a  little  proud  thereof."^^ 

We  thus  see  how  widely  the  different  races  of  man  differ  in 
their  taste  for  the  beautiful.  In  every  nation  sufficiently  advanced 
to  have  made  effigies  of  their  gods  or  of  their  deified  rulers,  the 
sculptors  no  doubt  have  endeavored  to  express  their  highest  ideal 
of  beauty  and  grandeur."  Under  this  point  of  view  it  is  vv^ell  to 
compare  in  our  mind  the  Jupiter  or  Apollo  of  the  Greeks  with  the 
Egyptian  or  Assyrian  statues;  and  these  with  the  hideous  bas- 
reliefs  on  the  ruined  buildings  of  Central  America. 

I  have  met  with  very  few  statements  opposed  to  this  conclusion. 
Mr.  Winwood  Reade,  however,  who  has  had  ample  opportunities 
for  observation,  not  only  with  the  negroes  of  the  West  Coast  of 
Africa,  but  with  those  of  the  interior  who  have  never  associated 
with  Europeans,  is  convinced  that  their  ideas  of  beauty  are  on 
the  whole  the  same  as  ours;  and  Dr.  Rohlfs  writes  to  me  to  the 
same  effect  with  respect  to  Bornu  and  the  countries  Inhabited  by 
the  Pulio  tribes.  Mr.  Reade  found  that  he  agreed  with  the  negroes 
in  their  estimation  of  the  beauty  of  the  native  girls;  and  that 
their  appreciation  of  the  beauty  of  European  women  corresponded 
with  ours.  They  admire  long  hair,  and  use  artificial  means  to 
make  it  appear  abundant;  they  admire  also  a  beard,  though 
themselves  very  scantily  provided.  Mr.  Reade  feels  doubtful  what 
kind  of  nose  is  most  appreciated:  a  girl  has  been  heard  to  say, 
"I  do  not  want  to  marry  him,  he  has  got  no  nose;"  and  this  shows 
that  a  very  flat  nose  is  not  admired.  We  should,  however,  bear 
in  mind  that  the  depressed,  broad  noses  and  projecting  jaws  of 
the  negroes  of  the  West  Coast  are  exceptional  types  with  the 
inhabitants  of  Africa.  Notwithstanding  the  foregoing  statements, 
Mr.  Reade  admits  that  negroes  "do  not  like  the,  color  of  our  skin; 
"they  look  on  blue  eyes  with  aversion,  and  they  think  our  noses 
"too  long  and  our  lips  too  thin."  He  does  not  think  it  probable 
that  negroes  would  ever  prefer  the  most  beautiful  European 
woman,  on  the  mere  grounds  of  physical  admiration,  to  a  good- 
looking  negress.^ 


<56  Dr.  Barnard  Davis  quotes  Mr.  Prichard  and  others  for  these  facts 
in  regard  to  the  Polynesians,  in  'Anthropological  Review,'  April,  1870, 
p.  185,  191. 

67  Ch.  Comte  has  remarks  to  this  effect  in  his  'Traite  de  Legislation,' 
3rd  edit.  1837,  p.  136. 

«8The  'African  Sketch  Book,'  vol.  ii.  1873,  pp.  253,  394,  521.  The 
Fuegians,  as  I  have  been  informed  by  a  missionary  who  long  resided 
with  them,  consider  European  women  as  extremely  beautiful;  but 
from  what  we  have  seen  of  the  judgment  of  the  other  aborigines  of 
America,  I  cannot  but  think  that  this  must  be  a  mistake,  unless  in- 
deed the  statement  refers  to  the  few  Fuegians  who  have  lived  for  some 
time  with  Europeans,  and  who  must  consider  us  as  superior  beings.    I 


MAN— BEAUTY.  577 

The  general  truth  of  the  principle,  long  ago  insisted  on  by 
Humboldt/^  that  man  admires  and  often  tries  to  exaggerate  what- 
ever characters  nature  may  have  given  him,  is  shown  in  many 
ways.  The  practice  of  beardless  races  extirpating  every  trace  of 
a  beard,  and  often  all  the  hairs  on  the  body,  affords  one  illustra- 
tion. The  skull  has  been  greatly  modified  during  ancient  and 
modern  times  by  many  nations;  and  there  can  be  little  doubt 
that  this  has  been  practiced,  especially  in  N.  and  S.  America,  in 
order  to  exaggerate  some  natural  and  admired  peculiarity.  Many 
American  Indians  are  known  to  admire  a  head  so  extremely  flat- 
tened as  to  appear  to  us  idiotic.  The  natives  on  the  north-western 
coast  compress  the  head  into  a  pointed  cone;  and  it  is  their  con- 
stant practice  to  gather  the  hair  into  a  knot  on  the  top  of  the 
head,  for  the  sake,  as  Dr.  Wilson  remarks,  "of  increasing  the  ap- 
"parent  elevation  of  the  favorite  conoid  form."  The  inhabitants 
of  Arakhan  "admire  a  broad,  smooth  forehead,  and  in  order  to  pro- 
"duce  it,  they  fasten  a  plate  of  lead  on  the  heads  of  the  new-born 
"children."  On  the  other  hand,  "a  broad,  well-rounded  occiput  is 
"considered  a  great  beauty"  by  the  natives  of  the  Fiji  islands.™ 

As  vath  the  skull,  so  with  the  nose;  the  ancient  Huns  during 
the  age  of  Attil-a  were  accustomed  to  flatten  the  noses  of  their 
infants  with  bandages,  "for  the  sake  of  exaggerating  a  natural 
"conformation."  With  the  Tahitians,  to  be  called  long-nose  is 
considered  an  insult,  and  they  compress  the  noses  and  fore- 
heads of  their  children  for  the  sake  of  beauty.  The  same  holds 
with  the  Malays  of  Sumatra,  the  Hottentots,  certain  Negroes, 
and  the  natives  of  Brazil.^^  The  Chinese  have  by  nature  un- 
usually small  feet;^^  and  it  is  well  known  that  the  women  of 
the  upper  classes  distort  their  feet  to  make  them  still  smaller. 
Lastly,  Humboldt  thinks  that  the  American  Indians  prefer  col- 


should  add  that  a  most  experienced  observer,  Capt.  Burton,  believes 
that  a  ■woman  whom  we  consider  beautiful  is  admired  throug-hout  the 
world,  'Anthropological  Review,'  March,  1864,  p.  245. 

69  'Personal  Narrative,'  Eng-.  translat.  vol.  iv.  p.  518,  and  elsewhere. 
Mantegazza,  in  his  'Yiaggi  e  Studi,'  1887,  strongly  insists  on  this  same 
principle. 

TO  On  the  skulls  of  the  American  tribes,  see  Nott  and  Gliddon,  'Types 
of  Mankind,'  1S54,  p.  440;  Prichard,  'Phys.  Hist,  of  Mankind,'  vol.  i. 
3rd  edit.  p.  321;  en  the  natives  of  Arakhan,  ibid.  vol.  iv.  p.  537.  Wilson, 
'Physical  Ethnology,'  Smithsonian  Institution,  1863,  p.  288;  on  the 
Fijians,  p.  290.  Sir  J.  Lubbock  ('Prehistoric  Times,'  2nd  edit.  1869,  p. 
506)  gives  an  excellent  resume  on  this  subject. 

Ti  On  the  Huns,  Godron,  'De  I'Espece,'  torn,  ii,  1859,  p.  SCO.  On  the 
Tahitians,  Waitz,  'Anthropolog.'  Eng.  translat.  vol.  1.  p.  305.  Marsden, 
quoted  by  Prichard,  'Phys.  Hist,  of  Mankind,'  3rd  edit.  vol.  v.  p.  67. 
Lawrence,   'Lectures  on  Physiology,'  p.  337. 

"  This  fact  was  ascertained  in  the  'Reise  der  Novara:    AnthropcZog. 
Theil,'  Dr.  Weisbach,  1867,  s.  285. 
3S 


578  THE  DESCENT  OF  MAN. 

oring  their  bodies  with  red  paint  in  order  to  exaggerate  their 
natural  tint;  and  until  recently  European  women  added  to  their 
naturally  bright  colors  by  rouge  and  white  cosmetics;  but  it  may 
be  doubted  whether  barbarous  nations  have  generally  had  any 
such  intention  in  painting  themselves. 

In  the  fashions  of  our  own  dress  we  see  exactly  the  same  prin- 
ciple and  the  same  desire  to  carry  every  point  to  an  extreme; 
we  exhibit,  also,  the  same  spirit  of  emulation.  But  the  fashions 
of  savages  are  far  more  permanent  than  ours;  and  whenever 
their  bodies  are  artificially  modified,  this  is  necessarily  the  case. 
The  Arab  women  of  the  Upper  Nile  occupy  about  three  days  in 
dressing  their  hair;  they  never  imitate  other  tribes,  "but  simply 
"vie  with  each  other  in  the  superlativeness  of  their  own  style." 
Dr.  Wilson,  in  speaking  of  the  compressed  skulls  of  various 
American  races,  adds,  "such  usages  are  among  the  least  eradi- 
"cable,  and  long  survive  the  shock  of  revolutions  that  change 
"dynasties  and  efface  more  important  national  peculiarities."^^ 
The  same  principle  comes  into  play  in  the  art  of  breeding;  and 
we  can  thus  understand,  as  I  have  elsewhere  explained,^*  the 
wonderful  development  of  the  many  races  of  animals  and  plants, 
which  have  been  kept  merely  for  ornament.  Fanciers  always 
wish  each  character  to  be  somewhat  increased;  they  do  not  ad- 
mire a  medium  standard;  they  certainly  do  not  desire  any  great 
and  abrupt  change  in  the  character  of  their  breeds;  they  admire 
solely  what  they  are  accustomed  to,  but  they  ardently  desire  to 
see  each  characteristic  feature  a  little  more  developed. 

The  senses  of  man  and  of  the  lower  animals  seem  to  be  so 
constituted  that  brilliant  colors  and  certain  forms,  as  well  as 
harmonious  and  rhythmical  sounds,  give  pleasure  and  are  called 
beautiful;  but  why  this  should  be  so,  we  know  not.  j  It  is  cer- 
tainly not  true  that  there  is  in  the  mind  of  man  anyliniversal 
standard  of  beauty  with  respect  to  the  human  body.  It  is,  how- 
ever, possible  that  certain  tastes  may  in  the  course  of  time  become 
inherited,  though  there  is  no  evidence  in  favor  of  this  belief;  and 
if  so,  each  race  would  possess  its  own  innate  ideal  standard  of 
beauty.  It  has  been  argued^^  that  ugliness  consists  in  an  approach 
to  the  structure  of  the  lower  animals,  and  no  doubt  this  is  partly 
true  with  the  more  civilized  nations,  in  which  intellect  is  highly 
appreciated;  but  this  explanation  will  hardly  apply  to  all  forms 
of  ugliness.  The  men  of  each  race  prefer  what  they  are  ac- 
customed to;    they  cannot  endure  any  great  change;    but  they 

'3  'Smithsonian  Institution,'  1863,  p.  289.  On  the  fashions  of  Arab 
women,  Sir  S.  Baker,  'The  Nile  Tributaries,'  1867,  p.  121. 

■^^  'The  "Variation  of  Animals  and  Plants  under  Domestication,'  vol.  i. 
p.  214;    vol.  ii,  p.  240. 

'5  Schaaffhausen,  'Archiv.  fur  Anthropologic,'  1866,  a  164. 


MAN— BEAQTT.  579 

like  variety,  and  admire  each  characteristic  carried  to  a  moderate 
extreme.'^  Men  accustomed  to  a  nearly  oval  face,  to  straight  and 
regular  features,  and  to  bright  colors,  admire,  as  we  Europeans 
know,  these  points  when  strongly  developed.  On  the  other  hand, 
men  accustomed  to  a  broad  face,  with  high  cheek-bones,  a  de- 
pressed nose,  and  a  black  skin,  admire  these  peculiarities  when 
strongly  marked.  No  doubt  characters  of  all  kinds  may  be  too 
much  developed  for  beauty.  Hence  a  perfect  beauty,  which  im- 
plies many  characters  modified  in  a  particular  manner,  will  be  in 
every  race  a  prodigy.  As  the  great  anatomist  Bichat  long  ago 
said,  if  every  one  were  cast  in  the  same  mold,  there  would  be  no 
such  thing  as  beauty.  If  all  our  women  v/ere  to  become  as  beau- 
tiful as  the  Venus  de  Medici,  we  should  for  a  time  be  charmed; 
but  we  should  soon  wish  for  variety;  and  as  soon  as  we  had 
obtained  variety,  we  should  wish  to  see  certain  characters  a  little 
exaggerated  beyond  the  then  existing  common  standard. 

•^s  Mr.  Bain  has  collected  ('Mental  and  Moral  Science,'  1868,  pp.  304-314) 
about  a  dozen  more  or  less  different  theories  of  the  idea  of  beauty; 
but  none  are  quite  the  same  as  that  here  given. 


580  THE  DESCENT  OF  MAN. 


CHAPTER   XX. 
SECONDARY  SEXUAL  CHARACTERS  OF  MAN— Continued. 

On  the  effects  of  the  continued  selection  of  women  according  to  a  dif- 
ferent standard  of  beauty  in  each  race— On  the  causes  which  inter- 
fere witli  sexual  selection  in  civilized  and  savage  nations— Condi- 
tions favorable  to  sexual  selection  during  primeval  times— On  the 
manner  of  action  of  sexual  selection  with  mankind— On  the  women 
in  savage  tribes  having  some  power  to  choose  their  husbands- 
Absence  of  hair  on  the  body,  and  development  of  the  beard— Color 
of  the  skin — Summary. 

We  have  seen  in  the  last  chapter  that  with  all  barbarous  races 
ornaments,  dress,  and  external  appearance  are  highly  valued; 
and  that  the  men  judge  of  the  beauty  of  their  women  by  widely 
different  standards.  We  must  next  inquire  whether  this  prefer- 
ence and  the  consequent  selection  during  many  generations  of 
those  v/omen,  which  appear  to  the  men  of  each  race  the  most 
attractive,  has  altered  the  character  either  of  the  females  alone, 
or  of  both  sexes.  With  mammals  the  general  rule  appears  to  be 
that  characters  of  all  kinds  are  inherited  equally  by  the  males 
and  females;  we  might  therefore  expect  that  with  mankind  any 
characters  gained  by  the  females  or  by  the  males  through  sexual 
selection,  would  commonly  be  transferred  to  the  offspring  of  both 
sexes.  If  any  change  has  thus  been  effected,  it  is  almost  certain 
that  the  different  races  would  be  differently  modified,  as  each  has 
its  own  standard  of  beauty. 

With  mankind,  especially  with  savages,  many  causes  interfere 
with  the  action  of  sexual  selection  as  far  as  the  bodily  frame  is 
concerned.  Civilized  men  are  largely  attracted  by  the  mental 
charms  of  women,  by  their  wealth,  and  especially  by  their  social 
position;  for  men  rarely  marry  into  a  much  lower  rank.  The 
men  who  succeed  in  obtaining  the  more  beautiful  women,  will 
not  have  a  better  chance  of  leaving  a  long  line  of  descendants 
than  other  men  with  plainer  wives,  save  the  few  who  bequeath 
their  fortunes  according  to  primogeniture.  With  respect  to  the 
opposite  form  of  selection,  namely  of  the  more  attractive  men  by 
the  women,  although  in  civilized  nations  women  have  free  cr 


MAN-SEXUAI^     SELECTION.  581 

almost  free  choice,  Y/liicIi  is  not  the  case  with  barbarous  races, 
yet  their  choice  is  largely  influenced  by  the  social  position  and 
wealth  of  the  men;  and  the  success  of  the  latter  in  life  depends 
much  on  their  intellectual  powers  and  energy,  or  on  the  fruits  of 
these  same  powers  in  their  forefathers.  No  excuse  vd  needed  for 
treating  this  subject  in  some  detail;  for,  as  the  German  philoso- 
pher Schopenhauer  remarks,  "the  final  aim  of  all  love  intrigues, 
"be  they  comic  or  tragic,  is  really  of  more  importance  than  all 
"other  ends  in  human  life.  What  it  all  turns  upon  is  nothing  less 
"than  the  composition  of  the  next  generation.  ...  It  is  not  the 
"weal  or  woe  of  any  one  individual,  but  that  of  the  human  race  to 
"come,  which  is  here  at  stake."^ 

There  is,  however,  reason  to  believe  that  in  certain  civilized 
and  semi-civilized  nations  sexual  selection  has  effected  soms- 
thing  in  modifying  the  bodily  frame  of  some  of  the  members. 
Many  persons  are  convinced,  as  it  appears  to  me  with  justice, 
that  our  aristocracy,  including  under  this  term  all  wealthy  fami- 
lies in  which  primogeniture  has  long  prevailed,  from  having 
chosen  during  many  generations  from  all  classes  the  more  beau- 
tiful women  as  their  wives,  have  become  handsomer,  according 
to  the  European  standard,  than  the  middle  classes;  yet  the 
middle  classes  are  placed  under  equally  favorable  conditions  of 
life  for  the  perfect  development  of  the  body.  Cook  remarks  that 
the  superiority  in  personal  appearance  "which  is  observable  in 
"the  erees  or  nobles  in  all  the  other  islands  (of  the  Pacific)  is 
"found  in  the  Sandwich  islands;"  but  this  may  be  chiefly  due 
to  their  better  food  and  manner  of  life. 

The  old  traveler  Chardin,  in  describing  the  Persians,  says  their 
*'blood  is  now  highly  refined  by  frequent  intermixtures  with  the 
"Georgians  and  Circassians,  two  nations  which  surpass  all  the 
"world  in  personal  beauty.  There  is  hardly  a  man  of  rank  in 
"Persia  who  is  not  born  of  a  Georgian  or  Circassian  mother."  He 
adds  that  they  inherit  their  beauty,  "not  from  their  ancestors, 
"for  v/ithout  the  above  mixture,  the  men  of  rank  in  Persia, 
"who  are  descendants  of  the  Tartars,  would  be  extremely  ugly."^ 
Here  is  a  more  curious  case;  the  priestesses  who  attended  the 
temple  of  Venus  Erycina  at  San-Giuliano  in  Sicily,  were  selected 
for  their  beauty  out  of  the  whole  of  Greece;  they  were  not  vestal 
virgins,  and  Quatrefages,^  v/ho  states  the  foregoing  fact,  says  that 
the  w^omen  of  San-Giuliano  are  now  famous  as  the  most  beautiful 


1  'Schopenhauer  and  Darwinism,'  in  'Journal  of  Anthropology,'  Jan. 
1871,  p.  323. 

"  These  quotations  are  taken  from  Lawrence  ('Lectures  on  Physiolo- 
gy,' &c.  1822,  p.  393),  who  attributes  the  beauty  of  the  upper  classes  in 
England  to  the  men  having  long  selected  the  more  beautiful  women. 

^  'Anthropologie,'  'Revue  des  Cours  Scientifiques,'  Oct.  1868,  p.  721. 

S8 


5S2  THE   DESCENT  OF  MAN. 

In  the  island,  and  are  sought  by  artists  as  models.     But  it  is 
obvious  that  the  evidence  in  all  the  above  cases  is  doubtful. 

The  following  case,  though  relating  to  savages,  is  well  worth 
giving  from  its  curiosity.  Mr.  Winwood  Reade  informs  me  that 
the  Jollofs,  a  tribe  of  negroes  on  the  west  coast  of  Africa,  "are 
"remarkable  for  their  uniformly  fine  appearance."  A  friend  of 
his  asked  one  of  these  men,  "How  is  it  that  every  one  whom  I 
"meet  is  so  fine-looking,  not  only  your  men,  but  your  women?" 
The  Jollof  answered,  "It  is  very  easily  explained:  it  has  always 
"been  our  custom  to  pick  out  our  worst-looking  slaves  and  to 
"sell  them."  It  need  hardly  be  added  that  with  all  savages, 
female  slaves  serve  as  concubines.  That  this  negro  should  have 
attributed,  whether  rightly  or  wrongly,  the  fine  appearance  of 
his  tribe  to  the  long-continued  elimination  of  the  ugly  women  is 
not  so  surprising  as  it  may  at  first  appear;  for  I  have  elsewhere 
shown*  that  negroes  fully  appreciate  the  importance  of  selection 
in  the  breeding  of  their  domestic  animals,  and  I  could  give  from 
Mr.  Reade  additional  evidence  on  this  head. 

The  Causes  which  prevent  or  check  the  Action  of  Sexual  SeleO' 
tion  with  Savages. — The  chief  causes  are,  first,  so-called  com- 
munal marriages  or  promiscuous  intercourse;  secondly,  the  con- 
sequences of  female  infanticide;  thirdly,  early  betrothals;  and 
lastly,  the  low  estimation  in  which  women  are  held,  as  mere 
slaves.    These  four  points  must  be  considered  in  some  detail. 

It  is  obvious  that  as  long  as  the  pairing  of  man,  or  of  any 
other  animal,  is  left  to  mere  chance,  with  nO'  choice  exerted  by 
either  sex,  there  can  be  no  sexual  selection;  and  no  effect  will  be 
produced  on  the  offspring  by  certain  individuals  having  had  an 
advantage  over  others  in  their  courtship.  Now  it  is  asserted  that 
there  exist  at  the  present  day  tribes  which  practice  what  Sir, 
J.  Lubbock  by  courtesy  calls  communal  marriages;  that  is,  all 
the  men  and  women  in  the  tribe  are  husbands  and  wives  to  one 
another.  The  licentiousness  of  many  savages  is  no  doubt  aston- 
ishing, but  it  seems  to  me  that  more  evidence  is  requisite,  before 
we  fully  admit  that  their  intercourse  is  in  any  case  promiscuous. 
Nevertheless  all  those  who  have  most  closely  studied  the  subject,^ 

*  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  i. 
p.  207. 

5  Sir  J.  Lubbock,  'The  Origin  of  Civilization,'  1870,  chap,  iii,  especially 
pp.  60-67.  Mr.  M'Lennan,  in  his  extremely  valuable  work  on  'Primitive 
Marriage,'  1865,  p.  163,  speaks  of  the  union  of  the  sexes  "in  the  earliest 
"times  as  loose,  transitory,  and  in  some  degree  promiscuous."  Mr. 
M'Lennan  and  Sir  J.  Lubbock  have  collected  much  evidence  on  the 
extreme  licentiousness  of  savages  at  the  present  time.  Mr.  L.  H. 
Morgan,  in  his  interesting  memoir  on  the  classificatory  system  of  re- 
lationship ('Proc.  American  Acad,  of  Sciences,'   vol.  vii.  Feb.  1868,  p. 


MAN— CHECKS  TO  SEXUAL  SELECTION.        583 

and  whose  judgment  is  worth  much  more  than  mine,  believe  that 
communal  marriage  (this  expression  being  variously  guarded) 
was  the  original  and  universal  form  throughout  the  world,  in- 
cluding therein  the  intermarriage  of  brothers  and  sisters.  The 
late  Sir  A.  Smith,  who  had  traveled  widely  in  S.  Africa,  and 
knew  much  about  the  habits  of  savages  there  and  elsewhere,  ex- 
pressed to  me  the  strongest  opinion  that  no  race  exists  in  which 
woman  is  considered  as  the  property  of  the  community.  I  believe 
that  his  judgment  was  largely  determined  by  what  is  implied  by 
the  term  marriage.  Throughout  the  following  discussion  I  use 
the  term  in  the  same  sense  as  when  naturalists  speak  of  animals 
as  monogamous,  meaning  thereby  that  the  male  is  accepted  by  or 
chooses  a  single  female,  and  lives  with  her  either  during  the 
breeding-season  or  for  the  whole  year,  keeping  possession  of 
her  by  the  law  of  might;  or,  as  when  they  speak  of  a  polygamous 
species,  meaning  that  the  male  lives  with  several  females.  This 
kind  of  marriage  is  all  that  concerns  us  here,  as  it  suffices  for 
the  work  of  sexual  selection.  But  I  know  that  some  of  the  writers 
above  referred  to,  imply  by  the  term  marriage,  a  recognized  right, 
protected  by  the  tribe. 

The  indirect  evidence  in  favor  of  the  belief  of  the  former  preva- 
lence of  communal  marriages  is  strong,  and  rests  chiefly  on  the 
terms  of  relationship  which  are  employed  between  the  members 
of  the  same  tribe,  implying  a  connection  with  the  tribe,  and  not 
with  either  parent.  But  the  subject  is  too  large  and  complex  for 
even  an  abstract  to  be  here  given,  and  I  will  confine  myself  to  a 
few  remarks.  It  is  evident  in  the  case  of  such  marriages,  or 
where  the  marriage  tie  is  very  loose,  that  the  relationship  of  the 
child  to  its  father  cannot  be  known.  But  it  seems  almost  incred- 
ible that  the  relationship  of  the  child  to  its  mother  should  ever 
be  completely  ignored,  especially  as  the  women  in  most  savage 
tribes  nurse  their  infants  for  a  long  time.  Accordingly,  in  many 
cases  the  lines  of  descent  are  traced  through  the  mother  alone, 
to  the  exclusion  of  the  father.  But  in  other  cases  the  terms  em- 
ployed express  a  connection  with  the  tribe  alone,  to  the  exclusion 
even  of  the  mother.  It  seems  possible  that  the  connection  be- 
tween the  related  members  of  the  same  barbarous  tribe,  exposed 
to  all  sorts  of  danger,  might  be  so  much  more  important,  owing 
to  the  need  of  mutual  protection  and  aid,  than  that  between  the 
mother  and  her  child,  as  to  lead  to  the  sole  use  of  terms  expres- 
sive of  the  former  relationships;  but  Mr.  Morgan  is  convinced 
that  this  view  is  by  no  means  sufficient. 


475),  concludes  that  polygamy  and  all  forms  of  marriage  during  pri- 
meval times  were  essentially  unknown.  It  appears  also,  from  Sir  J. 
Lubbock's  work,  that  Bachofen  likewise  believes  that  communal  inter* 
course  originally  prevailed. 


584  THE   DESCENT  OF  MAN. 

The  terms  of  relationsliip  used  in  different  parts  of  the  world 
may  be  divided,  according  to  the  author  just  quoted,  into  two 
great  classes,  the  classifi'catory  and  "descriptive, — the  latter  being 
employed  by  us.  It  is  the  classificatory  system  which  so  strongly 
leads  to  the  belief,  that  communal  and  other  extremely  loose 
forms  of  marriage  v/ere  originally  universal.  But  as  far  as  I  can 
see,  there  is  no  necessity  on  this  ground  for  believing  in  abso- 
lutely promiscuous  intercourse;  and  I  am  glad  to  find  that  this 
is  Sir  J.  Lubbock's  view.  Men  and  women,  like  many  of  the 
lower  animals,  might  formerly  have  entered  into  strict  though 
temporary  unions  for  each  birth,  and  in  this  case  nearly  as  much 
confusion  would  have  arisen  in  the  terms  of  relationship,  as  in 
the  case  of  promiscuous  intercourse.  As  far  as  sexual  selection 
is  concerned,  all  that  is  required  is  that  choice  should  be  exerted 
before  the  parents  unite,  and  it  signifies  little  whether  the  unions 
last  for  life  or  only  for  a  season. 

Besides  the  evidence  derived  from  the  terms  of  relationship, 
other  lines  of  reasoning  indicate  the  former  wide  prevalence  of 
communal  marriage.  Sir  J.  Lubbock  accounts''  for  the  strange 
and  widely-extended  habit  of  exogamy — that  is,  the  men  of  one 
tribe  taking  wives  from  a  distinct  tribe, — by  communism  having 
been  the  original  form  of  intercourse;  so  that  a  man  never  ob- 
tained a  v/ife  for  himself  unless  he  captured  her  from  a  neigh- 
boring and  hostile  tribe,  and  then  she  would  naturally  have  be- 
come his  sole  and  valuable  property.  Thus  the  practice  of  captur- 
ing wives  might  have  arisen;  and  from  the  honor  so  gained  it 
might  ultimately  have  become  the  universal  habit.  According 
to  Sir  J.  Lubbock,*'  we  can  also  thus  understand  "the  necessity 
"of  expiation  for  marriage  as  an  infringement  of  tribal  rites, 
"since,  according  to  old  ideas,  a  man  had  no  right  to  appropriate 
"to  himself  that  which  belonged  to  the  whole  tribe."  Sir  J.  Lub- 
bock further  gives  a  curious  body  of  facts  showing  that  in  old 
times  high  honor  was  bestowed  on  women  who  were  utterly  licen- 
tious; and  this,  as  he  explains,  is  intelligible,  if  we  admit  that 
promiscuous  intercourse  was  the  aboriginal,  and  therefore  long 
revered  custom  of  the  tribe.^ 

Although  the  manner  of  development  of  the  marriage-tie  is  an 
obscure  subject,  as  we  may  infer  from  the  divergent  opinions  on 
several  points  between  the  three  authors  who  have  studied  it 
most  closely,  namely,  Mr.  Morgan,  Mr.  M'Lennan,  and  Sir  J. 
Lubbock,  yet  from  the  foregoing  and  several  other  lines  of  evi- 


6  Address  to  British  Association  'On  the  Social  and  Religious  Con- 
dition of  the  Lower  Races  of  Man,'  1870,  p.  20. 

'^  'Origin  of  Civilization,'  1870,  p.  86.  In  the  several  works  above 
quoted,  there  will  be  found  copious  evidence  on  relationship  through 
the  females  alone,  or  with  the  tribe  alone. 


MAN— CHECKS  TO  SEXUAL  SELECTION.        585 

dence  it  seems  probable®  that  the  habit  of  marriage,  in  any  strict 
sense  of  the  word,  has  been  gradually  developed;     and  that  al- 
most promiscuous  or  very  loose  intercourse  was  once  extremely 
common  throughout  the  world.     Nevertheless  from  the.  strength 
of  the  feeling  of  jealousy  all  through  the  animal  kingdom,  as  well 
as  from  the  analogy  of  the  lower  animals,  more  particularly  of 
those  which  come  nearest  to  man,  I  cannot  believe  that  absolutely 
promiscuous  intercourse  prevailed  in  times  past,  shortly  before 
man  attained  to  his  present  rank  in  the  zoological  scale.    Man,  as 
I  have  attempted  to  show,  is  certainly  descended  from  some  ape- 
like creature.     With  the  existing  Quadrumana,  as  far  as  their 
habits  are  known,  the  males  of  some  species  are  monogamous, 
but  live  during  only  a  part  of  the  year  with  the  females;    of  this 
the  orang  seems  to  afford  an  instance.     Several  kinds,  for  ex- 
ample some  of  the  Indian  and  American  monkeys,  are  strictly 
monogamous,  and  associate  all  the  year  round  with  their  wives. 
Others  are  polygamous,  for  example  the  gorilla  and  several  Amer- 
ican species,  and  each  family  lives  separate.    Even  when  this  oc- 
curs, the  families  inhabiting  the  same  district  are  probably  some- 
what social:    the  chimpanzee,  for  instance,  is  occasionally  met 
with  in  large  bands.     Again,  other  species  are  polygamous,  but 
several  males,  each  with  his  own  females,  live  associated  in  a 
body,  as  with  several  species  of  baboons.^    We  may  indeed  con- 
clude from  what  we  know  of  the  jealousy  of  all  male  quadrupeds, 
armed,  as  many  of  them  are,  with  special  weapons  for  battling 
with  their  rivals,  that  promiscuous  intercourse  in  a  state  of  nature 
is  extremely  improbable.    The  pairing  may  not  last  for  life,  but 
only  for  each  birth;    yet  if  the  males  which  are  the  strongest  and 
best  able  to  defend  or  otherwise  assist  their  females  and  young, 
were  to  select  the  more  attractive  females,  this  would  suflace  for 
sexual  selection. 

Therefore,  looking  far  enough  back  in  the  stream  of  time,  and 
judging  from  the  social  habits  of  man  as  he  now  exists,  the  most 
probable  view  is  that  he  aboriginally  lived  in  small  communities, 
each  with  a  single  wife,  or  if  powerful  with  several,  whom  he 
jealously  guarded  against  all  other  men.  Or  he  may  not  have 
been  a  social  animal,  and  yet  have  lived  with  several  wives,  like 
the  gorilla;    for  all  the  natives  "agree  that  but  one  adult  male 

s  Mr.  C.  Staniland  Wake  argues  strongly  ('Anthropologia,'  March, 
1874,  p.  197)  against  the  views  held  by  these  three  writers  on  the  former 
prevalence  of  almost  promiscuous  intercourse;  and  he  thinks  that  the 
classificatory  system  of  relationship  can  be  otherwise  explained. 

» Brehm  ('Illust.  Theirleben,'  B.  i.  p.  77)  says  Cjmocephalus  hama- 
dryas  lives  in  great  troops  containing  twice  as  many  adult  females  as 
adult  males.  See  Rengger  on  American  polygamous  species,  and  Owen 
('Anat.  of  Vertebrates,'  vol.  iii.  p.  746)  on  American  monogamous  spe- 
cies.   Other  references  might  be  added. 


586  THE  DESCENT  OF  MAN. 

"is  seen  in  a  band;  when  the  young  male  grows  up,  a  contest 
"takes  place  for  mastery,  and  the  strongest,  by  killing  and  driv- 
"ing  out  the  others,  establishes  himself  as  the  head  of  the  com- 
"munity."^"  The  younger  males,  being  thus  expelled  and  wan- 
dering about,  would,  when  at  last  successful  in  finding  a  partner, 
prevent  too  close  interbreeding  within  the  limits  of  the  same 
family. 

Although  savages  are  now  extremely  licentious,  and  although 
communal  marriages  may  formerly  have  largely  prevailed,  yet 
many  tribes  practice  some  form  of  marriage,  but  of  a  far  more  lax 
nature  than  that  of  civilized  nations.  Polygamy,  as  just  stated, 
is  almost  universally  followed  by  the  leading  men  in  every  tribe. 
Nevertheless  there  are  tribes,  standing  almost  at  the  bottom  of 
the  scale,  which  are  strictly  monogamous.  This  is  the  case  with 
the  Veddahs  of  Ceylon:  they  have  a  saying,  according  to  Sir  J. 
Lubbock,"  "that  death  alone  can  separate  husband  and  wife." 
An  intelligent  Kandyan  chief,  of  course  a  polygamist,  "was  per- 
"fectly  scandalized  at  the  utter  barbarism  of  living  with  only 
"one  wife,  and  never  parting  until  separated  by  death."  It  was, 
he  said,  "Just  like  the  Wanderoo  monkeys."  Whether  savages 
who  now  enter  into  some  form  of  marriage,  either  polygamous  or 
monogamous,  have  retained  this  habit  from  primeval  times,  or 
whether  they  have  returned  to  some  form  of  marriage,  after  pass- 
ing through  a  stage  of  promiscuous  intercourse,  I  will  not  pretend 
to  conjecture. 

Infanticide.— This  practice  is  now  very  common  throughout  the 
world,  and  there  is  reason  to  believe  that  it  prevailed  much  more 
extensively  during  former  times.^^  Barbarians  find  it  difiicult  to 
support  themselves  and  their  children,  and  it  is  a  simple  plan  to 
kill  their  infants.  In  South  America  some  tribes,  according  to 
Azara,  formerly  destroyed  so  many  infants  of  both  sexes,  that 
they  were  on  the  point  of  extinction.  In  the  Polynesian  Islands 
women  have  been  known  to  kill  from  four  or  five  to  even  ten  of 
their  children;  and  Ellis  could  not  find  a  single  woman  who  had 
not  killed  at  least  one.  Wherever  infanticide  prevails  the  strug- 
gle for  existence  will  be  in  so  far  less  severe,  and  all  the  members 
of  the  tribe  will  have  an  almost  equally  good  chance  of  rearing 
their  few  surviving  children.  In  most  cases  a  larger  number  of 
female  than  of  male  infants  are  destroyed,  for  it  is  obvious  that  the 
latter  are  of  more  value  to  the  tribe,  as  they  will,  when  grown  up, 
aid  in  defending  it,  and  can  support  themselves.    But  the  trouble 

10  Dr.  Savage,  in  'Boston  Journal  of  Nat.  Hist.'  vol.  v.  1845-47,  p.  423. 

11  'Prehistoric  Times,'  1869,  p.  424. 

1-  Mr.  M'Lennan,   'Primitive  Marriage,'  1865.    See  especially  on  exo- 
gamy and  infanticide,  pp.  130.  ASS.  3^. 


MAN-CHECKS  TO  SEXUAL  SELECTION.  587 

experienced  by  the  women  in  rearing  children,  their  consequent 
loss  of  beauty,  the  higher  estimation  set  on  them  when  few  and 
their  happier  fate,  are  assigned  by  the  women  themselves,  and 
by  various  observers,  as  additional  motives  for  infanticide.  In 
Australia,  where  female  infanticide  is  still  common.  Sir  G.  Grey 
estimated  the  proportion  of  native  women  to  men  as  one  to  three; 
but  others  say  as  two  to  three.  In  a  village  on  the  eastern  frontier 
of  India,  Colonel  MacCulloch  found  not  a  single  female  child.^^ 

When,  owing  to  female  infanticide,  the  women  of  a  tribe  were 
few,  the  habit  of  capturing  wives  from  neighboring  tribes  would 
naturally  arise.  Sir  J.  Lubbock,  however,  as  we  have  seen,  at- 
tributes the  practice  in  chief  part,  to  the  former  existence  of  com- 
munal marriage,  and  to  the  men  having  consequently  captured 
women  from  other  tribes  to  hold  as  their  sole  property.  Addi- 
tional causes  might  be  assigned,  such  as  the  communities  being 
very  small,  in  which  case,  marriageable  women  would  often  be 
deficient.  That  the  habit  was  most  extensively  practiced  during 
former  times,  even  by  the  ancestors  of  civilized  nations,  is  clearly 
shown  by  the  preservation  of  many  curious  customs  and  cere- 
monies, of  which  Mr.  M'Lennan  has  given  an  interesting  account. 
In  our  own  marriages  the  "best  man"  seems  originally  to  have 
been  the  chief  abettor  of  the  bridegroom  in  the  act  of  capture. 
Now  as  long  as  men  habitually  procured  their  wives  through  vio- 
lence and  craft,  they  would  have  been  glad  to  seize  on  any  wom- 
an, and  would  not  have  selected  the  more  attractive  ones.  But 
as  soon  as  the  practice  of  procuring  wives  from  a  distant  tribe 
was  effected  through  barter,  as  now  occurs  in  many  places,  the 
more  attractive  women  would  generally  have  been  purchased. 
The  Incessant  crossing,  however,  between  tribe  and  tribe,  which 
necessarily  follows  from  any  form  of  this  habit,  would  tend  to 
keep  all  the  people  inhabiting  the  same  country  nearly  uniform 
in  character;  and  this  would  interfere  with  the  power  of  sexual 
selection  in  differentiating  the  tribes. 

The  scarcity  of  women,  consequent  on  female  infanticide,  leads, 
also,  to  another  practice,  that  of  polyandry,  still  common  in  sev- 
eral parts  of  the  world,  and  which  formerly,  as  Mr.  M'Lennan 
believes,  prevailed  almost  universally;  but  this  latter  conclusion 
is  doubted  by  Mr.  Morgan  and  Sir  J.  Lubbock."  Whenever  two 
or  more  men  are  compelled  to  marry  one  woman,  it  is  certain 

13  Dr.  Gerland  ('Ueber  das  Aussterben  der  Naturvolker,'  1868)  has  col- 
lected much  information  on  infanticide,  see  especially  s.  27,  51,  54. 
Azara  ('Voyag-es,'  &c.  tom.  ii.  pp.  94,  116)  enters  in  detail  on  the  mo- 
tives.   See  also  M'Lennan  (ibid.  p.  139)  for  cases  in  India. 

"  'Primitive  Marriage,'  p.  208;  Sir  J.  Lubbock,  'Origin  of  Civiliza- 
tion,' p.  100.  See  also  Mr.  Morgan,  loo.  cit.,  on  the  former  prevalence 
of  polyandry. 


588  THE  DESCENT  OP  MAN. 

that  all  the  women  of  the  tribe  will  get  married,  and  there  will 
be  no  selection  by  the  men  of  the  more  attractive  women.  But 
under  these  circumstances  the  women  no  doubt  will  have  the 
power  of  choice,  and  will  prefer  the  more  attractive  men.  Azara, 
for  instance,  describes  how  carefully  a  Guana  woman  bargains 
for  all  sorts  of  privileges,  before  accepting  some  one  or  more  hus- 
bands; and  the  men  in  consequence  take  unusual  care  of  their  per- 
sonal appearance.  So  amongst  the  Todas  of  India,  who  practice 
polyandry,  the  girls  can  accept  or  refuse  any  man.^^  A  very  ugly 
man  in  these  cases  would  perhaps  altogether  fail  in  getting  a  wife, 
or  get  one  later  in  life;  but  the  handsomer  men,  although  more 
successful  in  obtaining  wives,  would  not,  as  far  as  we  can  see, 
leave  more  offspring  to  inherit  their  beauty  than  the  less  hand- 
somer husbands  of  the  same  women. 

Early  Betrothals  and  Slavery  of  Women.  — ^With  many  savages 
it  is  the  custom  to  betroth  the  females  whilst  mere  infants;  and 
this  would  effectually  prevent  preference  being  exerted  on  either 
side  according  to  personal  appearance.  But  it  would  not  prevent 
the  more  attractive  women  from  being  afterwards  stolen  or  taken 
by  force  from  their  husbands  by  the  more  powerful  men;  and 
this  often  happens  in  Australia,  America,  and  elsewhere.  The 
same  consequences  with  reference  to  sexual  selection  would  to  a 
certain  extent  follow,  when  women  are  valued  almost  solely  as 
slaves  or  beasts  of  burden,  as  is  the  case  with  many  savages. 
The  men,  however,  at  all  times  would  prefer  the  handsomest 
slaves  according  to  their  standard  of  beauty. 

We  thus  see  that  several  customs  prevail  with  savages  which 
must  greatly  interfere  with,  or  completely  stop,  the  action  of 
sexual  selection.  On  the  other  hand,  the  conditions  of  life  to 
which  savages  are  exposed,  and  some  of  their  habits,  are  favor- 
able to  natural  selection;  and  this  comes  into  play  at  the  same 
time  with  sexual  selection.  Savages  are  known  to  suffer  severely 
from  recurrent  famines;  they  do  not  increase  their  food  by  artifi- 
cial means;  they  rarely  refrain  from  marriage,^"  and  generally 
marry  whilst  young.  Consequently  they  must  be  subjected  to 
occasional  hard  struggles  for  existence,  and  the  favored  individ' 
uals  will  alone  survive. 

At  a  very  early  period,  before  man.  attained  to  his  present 
rank  in  the  scale,  many  of  his  conditions  would  be  different  from 

15  Azara,  'Voyag-es,'  &c.  torn.  ii.  pp.  92-95.  Colonel  Marshall,  'Amongst 
the  Todas,'  p.  212. 

isBurchell  says  ('Travels  in  S.  Africa,'  vol.  ii.  1824,  p.  58),  that 
among  the  wild  nations  of  Southern  Africa,  neither  men  nor  women 
ever  pass  their  lives  in  a  state  of  celibacy.  Azara  ('Voyag-es  dans 
I'Amerique  Merid.'  torn.  ii.  1809,  p.  21)  makes  precisely  the  same  re- 
mark in  regard  to  the  wild  Indians  of  South  America. 


MAN— MODE  OF  SEXUAL  SELECTION.  589 

what  now  obtains  amongst  savages.  Judging  from  the  analogy 
of  the  lower  animals  he  would  then  either  live  with  a  single 
female,  or  be  a  polygamist.  The  most  powerful  and  able  males 
would  succeed  best  in  obtaining  attractive  females.  They  would 
also  succeed  best  in  the  general  struggle  for  life,  and  in  defend- 
ing their  females,  as  well  as  their  offspring,  from  enemies  of  all 
kinds.  At  this  early  period  the  ancestors  of  man  would  not  be 
sufficiently  advanced  in  intellect  to  look  forward  to  distant  con- 
tingencies; they  would  not  foresee  that  the  rearing  of  all  their 
children,  especially  their  female  children,  would  make  the  struggle 
for  life  severer  for  the  tribe.  They  would  be  governed  more  by 
their  instincts  and  less  by  their  reason,  than  are  savages  at  the 
present  day.  They  would  not  at  that  period  have  partially  lost 
one  of  the  strongest  of  all  instincts,  common  to  all  the  lower 
animals,  namely  the  love  of  their  young  offspring;  and  conse- 
quently they  would  not  have  practiced  female  infanticide.  Wom- 
en would  not  have  been  thus  rendered  scarce,  and  polyandry 
would  not  have  been  practiced;  for  hardly  any  other  cause,  ex- 
cept the  scarcity  of  women  seems  sufficient  to  break  down  the 
natural  and  widely  prevalent  feeling  of  jealousy,  and  the  desire  of 
each  male  to  possess  a  female  for  himself.  Polyandry  would  be  a 
natural  stepping-stone  to  communal  marriages  or  almost  promis- 
cuous intercourse;  though  the  best  authorities  believe  that  this 
latter  habit  preceded  polyandry.  During  primordial  times  there 
would  be  no  early  betrothals,  for  this  implies  foresight.  Nor 
would  women  be  valued  merely  as  useful  slaves  or  beasts  of  bur- 
then. Both  sexes,  if  the  females  as  well  as  the  males  were  per- 
mitted to  exert  any  choice,  would  choose  their  partners  not  for 
mental  charms,  or  property,  or  social  position,  but  almost  solely 
from  external  appearance.  All  the  adults  would  marry  or  pair, 
and  all  the  offspring,  as  far  as  that  was  possible,  would  be  reared; 
so  that  the  struggle  for  existence  would  be  periodically  exces- 
sively severe.  Thus  during  these  times  all  the  conditions  for  sex- 
ual selection  would  have  been  more  favorable  than  at  a  later 
period,  when  man  had  advanced  in  his  intellectual  powers  but 
had  retrograded  in  his  instincts.  Therefore,  whatever  influence 
sexual  selection  may  have  had  in  producing  the  differences  be- 
tween the  races  of  man,  and  between  man  and  the  higher  Quad- 
rumana,  this  influence  would  have  been  more  powerful  at  a  re- 
mote period  than  at  the  present  day,  though  probably  not  yet 
wholly  lost. 

The  Manner  of  Action  of  Sexual  Selection  with  Mankind.  — With 
primeval  men  under  the  favorable  conditions  just  stated,  and 
with  those  savages  who  at  the  present  time  enter  into  any  mar- 
riage tie,  sexual  selection  has  probably  acted  in  the  following 
manner,  subject  to  greater  or  less  interference  from  female  iw 


590  THE  DESCENT  Oli:  MAN. 

fanticide,  early  betrothals,  &c.  The  strongest  and  most  vigorous 
men, — those  who  could  best  defend  and  hunt  for  their  families, 
who  were  provided  with  the  best  weapons  and  possessed  the 
most  property,  such  as  a  large  number  of  dogs  or  other  animals, — 
would  succeed  in  rearing  a  greater  average  number  of  offspring 
than  the  weaker  and  poorer  members  of  the  same  tribes.  There 
can,  also,  be  no  doubt  that  such  men  would  generally  be  able  to 
select  the  more  attractive  women.  At  present  the  chief?  of  nearly 
every  tribe  throughout  the  world  succeed  in  obtaining  more  than 
one  wife.  I  hear  from  Mr.  Mantell,  that  until  recently,  almost 
every  girl  in  New  Zealand,  who  was  pretty,  or  promised  to  be 
pretty,  was  tapu  to  some  chief.  With  the  Kafirs,  as  Mr.  C.  Hamil- 
ton states,^^  "the  chiefs  generally  have  the  pick  of  the  women  for 
"many  miles  round,  and  are  most  persevering  in  establishing  or 
"confirming  their  privilege."  We  have  seen  that  each  race  has 
its  own  style  of  beauty,  and  we  know  that  it  is  natural  to  man  to 
admire  each  characteristic  point  in  his  domestic  animals,  dress, 
ornaments,  and  personal  appearance,  when  carried  a  little  be- 
yond the  average.  If  then  the  several  foregoing  propositions  be 
admitted,  and  I  cannot  see  that  they  are  doubtful,  it  would  be  an 
inexplicable  circumstance,  if  the  selection  of  the  more  attractive 
women  by  the  more  powerful  men  of  each  tribe,  who  would  rear  on 
an  average  a  greater  number  of  children,  did  not  after  the  lapse 
of  many  generations  somewhat  modify  the  character  of  the  tribe. 
When  a  foreign  breed  of  our  domestic  animals  is  introduced 
into  a  new  country,  or  when  a  native  breed  is  long  and  carefully 
attended  to,  either  for  use  or  ornament,  it  is  found  after  several 
generations  to  have  undergone  a  greater  or  less  amount  of  change, 
whenever  the  means  of  comparison  exist.  This  follows  from  un- 
conscious selection  during  a  long  series  of  generations — that  is, 
the  preservation  of  the  most  approved  individuals — ^without  any 
wish  or  expectation  of  such  a  result  on  the  part  of  the  breeder. 
So  again,  if  during  many  years  two  careful  breeders  rear  animals 
of  the  same  family,  and  do  not  compare  them  together  or  with  a 
common  standard,  the  animals  are  found  to  have  become,  to  the 
surprise  of  their  owners,  slightly  different.^^  Each  breeder  has 
impressed,  as  Von  Nathusius  well  expresses  it,  the  character  of 
his  own  mind — his  own  taste  and  judgment — on  his  animals. 
What  reason,  then,  can  be  assigned  why  similar  results  should 
not  follow  from  the  long-continued  selection  of  the  most  admired 
women  by  those  men  of  each  tribe,  who  were  able  to  rear  the 
greatest  number  of  children?  This  would  be  unconscious  selec- 
tion, for  an  effect  would  be  produced,  independently  of  any  wish 

1'^  'Anthropological  Review,'  Jan.  1870,  p,  xvi. 

18  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
pp.  210-217. 


MAN— MODE  OF  SEXUAL  SELECTION.  591 

or  expectation  on  the  part  of  tlie  men  who  preferred  certain  women 
to  others. 

Let  us  suppose  the  members  of  a  tribe,  practicing  some  form 
of  marriage,  to  spread  over  an  unoccupied  continent;  they  would 
soon  split  up  into  distinct  hordes,  separated  from  each  other  by 
various  barriers,  and  still  more  effectually  by  the  incessant  wars 
between  all  barbarous  nations.  The  hordes  would  thus  be  ex- 
posed to  slightly  different  conditions  and  habits  of  life,  and  would 
sooner  or  later  come  to  differ  in  some  small  degree.  As  soon 
as  this  occurred,  each  isolated  tribe  would  form  for  itself  a  slight- 
ly different  standard  of  beauty; ^^  and  then  unconscious  selection 
would  come  into  action  through  the  more  powerful  and  leading 
men  preferring  certain  women  to  others.  Thus  the  differences  be- 
tween the  tribes,  at  first  very  slight,  would  gradually  and  inevi- 
tably be  more  or  less  increased. 

With  animals  in  a  state  of  nature,  many  characters  proper  to 
the  males,  such  as  size,  strength,  special  weapons,  courage  and 
pugnacity,  have  been  acquired  through  the  law  of  battle.  The 
semi-human  progenitors  of  man,  like  their  allies  the  Quadrumana, 
will  almost  certainly  have  been  thus  modified;  and,  as  savages 
still  fight  for  the  possession  of  their  women,  a  similar  process  of 
selection  has  probably  gone  on  in  a  greater  or  less  degree  to  the 
present  day.  Other  characters  proper  to  the  males  of  the  lower 
animals,  such  as  bright  colors  and  various  ornaments,  have  been 
acquired  by  the  more  attractive  males  having  been  preferred  by 
the  females.  There  are,  however,  exceptional  cases  in  which  the 
males  are  the  selecters,  instead  of  having  been  the  selected.  We 
recognize  such  cases  by  the  females  being  more  highly  orna- 
mented than  the  males, — their  ornamental  characters  having  been 
transmitted  exclusively  or  chiefly  to  their  female  offspring.  One 
such  case  has  been  described  in  the  order  to  which  man  belongs, 
that  of  the  Rhesus  monkey. 

Man  is  more  powerful  in  body  and  mind  than  woman,  and  in 
the  savage  state  he  keeps  her  in  a  far  more  abject  state  of  bond- 
age, than  does  the  male  of  any  other  animal;  therefore  it  is  not 
surprising  that  he  should  have  gained  the  power  of  selection. 
Women  are  everywhere  conscious  of  the  value  of  their  own  beauty; 
and  when  they  have  the  means,  they  take  more  delight  in  decorat- 
ing themselves  with  all  sorts  of  ornaments  than  do  men.  They 
borrow  the  plumes  of  male  birds,  with  which  nature  has  decked 
this  sex  in  order  to  charm  the  females.    As  women  have  long  been 

"  An  ing-enious  writer  argrues,  from  a  comparison  of  the  pictures  of 
Raphael,  Rubens,  and  modern  French  artists,  that  the  idea  of  beauty 
is  not  absolutely  the  same  even  throug-hout  Europe:  see  the  'Lives 
of  Haydn  and  Mozart,'  by  Bombet  (otherwise  M.  Beyle),  English  trans- 
lat.  p.  278. 


692  THE  DESCENT  OF  MAN. 

selected  for  beauty,  it  is  not  surprising  that  some  of  their  succes- 
sive variations  should  have  been  transmitted  exclusively  to  the 
same  sex;  consequently  that  they  should  have  transmitted  beauty 
in  a  somewhat  higher  degree  to  their  female  than  to  their  male 
offspring,  and  thus  have  become  more  beautiful,  according  to 
general  opinion,  than  men.  Women,  however,  certainly  transmit 
most  of  their  characters,  including  some  beauty,  to  their  offspring 
of  both  sexes;  so  that  the  continued  preference  by  the  men  of 
each  race  for  the  more  attractive  vfomen,  according  to  their  stand- 
ard of  taste,  will  have  tended  to  modify  in  the  same  manner  all 
the  individuals  of  both  sexes  belonging  to  the  race. 

With  respect  to  the  other  form  of  sexual  selection  (which  with 
the  lower  animals  is  much  the  more  common),  namely,  when  the 
females  are  the  selecters,  and  accept  only  those  males  which 
excite  or  charm  them  most,  we  have  reason  to  believe  that  it 
formerly  acted  on  our  progenitors.  Man  in  all  probability  owes 
his  beard,  and  perhaps  some  other  characters,  to  inheritance  from 
an  ancient  progenitor  who  thus  gained  his  ornaments.  But  this 
form  of  selection  may  have  occasionally  acted  during  later  times; 
for  in  utterly  barbarous  tribes  the  women  have  more  power  in 
choosing,  rejecting,  and  tempting  their  lovers,  or  of  afterwards 
changing  their  husbands,  than  might  have  been  expected.  As 
this  is  a  point  of  some  importance,  I  will  give  in  detail  such  evi- 
dence as  I  have  been  able  to  collect. 

Hearne  describes  how  a  woman  in  one  of  the  tribes  of  Arctic 
America  repeatedly  ran  away  from  her  husband  and  joined  her 
lover;  and  with  the  Charruas  of  S.  America,  according  to  Azara, 
divorce  is  quite  optional.  Amongst  the  Abipones,  a  man  on 
choosing  a  wife,  bargains  with  the  parents  about  the  price.  But 
"it  frequently  happens  that  the  girl  rescinds  what  has  been 
"agreed  upon  between  the  parents  and  the  bridegroom,  obstinate- 
"ly  rejecting  the  very  mention  of  marriage."  She  often  runs 
away,  hides  herself,  and  thus  eludes  the  bridegroom.  Captain 
Musters,  who  lived  with  the  Patagonians,  says  that  their  mar- 
riages are  always  settled  by  inclination;  "if  the  parents  make  a 
"match  contrary  to  the  daughter's  will,  she  refuses  and  is  never 
"compelled  to  comply."  In  Tierra  del  Fuego  a  young  man  first 
obtains  the  consent  of  the  parents  by  doing  them  some  service, 
and  then  he  attempts  to  carry  off  the  girl;  "but  if  she  is  unwilling, 
"she  hides  herself  in  the  woods  until  her  admirer  is  heartily  tired 
"of  looking  for  her,  and  gives  up  the  pursuit;  but  this  seldom 
"happens."  In  the  Fiji  Islands  the  man  seizes  on  the  woman 
whom  he  wishes  for  his  wife  by  actual  or  pretended  force;  but 
"on  reaching  the  home  of  her  abductor,  should  she  not  approve 
"of  the  match,  she  runs  to  some  one  who  can  protect  her;  if, 
"however,  she  is  satisfied,  the  matter  is  settled  forthwith."  With 
the  Kalmucks  there  is  a  regular  race  between  the  bride  and  bride- 


Man— MODE  OF  sexual  selection.  593 

groom,  tlie  former  having  a  fair  start;  and  Clarke  "was  assured 
"that  no  instance  occurs  of  a  girl  being  caught,  unless  she  has  a 
"partiality  to  the  pursuer."  Amongst  the  wild  tribes  of  the  Malay- 
Archipelago  there  is  also  a  racing  match;  and  it  appears  from  M. 
Bourien's  account,  as  Sir  J.  Lubbock  remarks,  that  "the  race  'is 
"  'not  to  the  swift,  nor  the  battle  to  the  strong,'  but  to  the  young 
"man  who  has  the  good  fortune  to  please  his  intended  bride."  A 
similar  custom,  with  the  same  result,  prevails  with  the  Koraks 
of  North-Eastern  Asia. 

Turning  to  Africa:  the  Kafirs  buy  their  wives,  and  girls  are 
severely  beaten  by  their  fathers  if  they  will  not  accept  a  chosen 
husband;  but  it  is  manifest  from  many  facts  given  by  the  Rev. 
Mr.  Shooter,  that  they  have  considerable  power  of  choice.  Thus 
very  ugly,  though  rich  men,  have  been  known  to  fail  in  getting 
wives.  The  girls,  before  consenting  to  be  betrothed,  compel  the 
men  to  show  themselves  off  first  in  front  and  then  behind,  and 
"exhibit  their  paces."  They  have  been  known  to  propose  to  a 
man,  and  they  not  rarely  run  away  with  a  favored  lover.  So 
again,  Mr.  Leslie,  who  was  intimately  acquainted  with  the  Kafirs, 
says,  "it  is  a  mistake  to  imagine  that  a  girl  is  sold  by  her  father 
"in  the  same  manner,  and  with  the  same  authority,  with  which 
"he  would  dispose  of  a  cow."  Amongst  the  degraded  Bushmen  of 
S.  Africa,  "when  a  girl  has  grown  up  to  womanhood  without  hav- 
"ing  been  betrothed,  which,  however,  does  not  often  happen,  her 
"lover  must  gain  her  approbation,  as  well  as  that  of  the  par- 
"ents."^°  Mr.  Winwood  Reade  made  inquiries  for  me  with  respect 
to  the  negroes  of  Western  Africa,  and  he  informs  me  that  "the 
"women,  at  least  among  the  more  intelligent  Pagan  tribes,  have 
"no  difficulty  in  getting  the  husbands  whom  they  may  desire, 
"although  it  is  considered  unwomanly  to  ask  a  man  to  marry 
"them.  They  are  quite  capable  of  falling  in  love,  and  of  forming 
"tender,  passionate,  and  faithful  attachments."  Additional  cases 
could  be  given. 

We  thus  see  that  with  savages  the  women  are  not  in  quite  so 
abject  a  state  in  relation  to  marriage,  as  has  often  been  supposed. 
They  can  tempt  the  men  whom  they  prefer,  and  can  sometimes 

20  Azara,  'Voyag-es,'  &c.  torn.  ii.  p.  23.  Dobrizhoffer,  'An  account  of 
the  Abipones,'  vol.  ii.  1822,  p.  207.  Capt.  Musters,  in  'Proc.  R.  Geo- 
graph.  Soc.,'  vol.  xv.  p.  47.  Willianis  on  the  Fiji  Islanders,  as  quoted 
by  Lubbock,  'Origin  of  Civilization,'  1870,  p.  79,  On  the  Puegians, 
King  and  FitzRoy,  'Voyages  of  the  Adventure  and  Beagle,'  vol.  ii. 
1839,  p.  182.  On  the  Kalmucks,  quoted  by  M'Lennan,  'Primitive  Mar- 
riage,' 1865,  p.  32.  On  the  Malays,  Lubbock,  ibid.  p.  76.  The  Rev.  J. 
Shooter,  'On  the  Kafirs  of  Natal,'  1S57,  pp.  52-60.  Mr.  D.  Leslie,  'Kafir 
Character  and  Customs,'  1871,  p.  4.  On  the  Bush-men,  Burchell,  'Trav- 
els in  S.  Africa,'  vol.  ii.  1824,  p.  59,  On  the  Koraks  by  McKennan,  as 
quoted  by  Mr.  Wake,  in  'Anthropologia,'  Oct.  1873,  p.  75. 
89 


594  THE  DESCENT  OF  MAN. 

reject  those  whom  they  dislike,  either  before  or  after  marriage. 
Preference  on  the  part  of  the  women,  steadily  acting  in  any  one 
direction,  would  ultimately  affect  the  character  of  the  tribe;  for 
the  women  would  generally  choose  not  merely  the  handsomest 
men,  according  to  their  standard  of  taste,  but  those  who  were  at 
the  same  time  best  able  to  defend  and  support  them.  Such  well- 
endowed  pairs  would  commonly  rear  a  larger  number  of  offspring 
than  the  less  favored.  The  same  result  would  obviously  follow 
in  a  still  more  marked  manner,  if  there  was  selection  on  both 
sides;  that  is  if  the  more  attractive,  and  at  the  same  time  more 
powerful  men  were  to  prefer,  and  were  preferred  by,  the  more  at- 
tractive women.  And  this  double  form  of  selection  seems  actually 
to  have  occurred,  especially  during  the  earlier  periods  of  our  long 
history. 

We  will  now  examine  a  little  more  closely  some  of  the  char- 
acters which  distinguish  the  several  races  of  man  from  one  an- 
other and  from  the  lower  animals,  namely,  the  greater  or  less 
deficiency  of  hair  on  the  body,  and  the  color  of  the  skin.  We 
need  say  nothing  about  the  great  diversity  in  the  shape  of  the 
features  and  of  the  skull  between  the  different  races,  as  we  have 
seen  in  the  last  chapter  how  different  is  the  standard  of  beauty  in 
these  respects.  These  characters  will  therefore  probably  have 
been  acted  on  through  sexual  selection;  but  we  have  no  means  of 
judging  whether  they  have  been  acted  on  chiefly  from  the  male 
or  female  side.  The  musical  faculties  of  man  have  likewise  been 
already  discussed. 

Absence  of  Hair  on  the  Body,  and  its  Development  on  the  Face 
and  Head. — From  the  presence  of  the  woolly  hair  or  lanugo  on 
the  human  foetus,  and  of  rudimentary  hairs  scattered  over  the 
body  during  maturity,  we  may  infer  that  man  is  descended  from 
some  animal  which  was  born  hairy  and  remained  so  during  life. 
The  loss  of  hair  is  an  inconvenience  and  probably  an  injury  to 
man,  even  in  a  hot  climate,  for  he  is  thus  exposed  to  the  scorch- 
ing of  the  sun,  and  to  sudden  chills,  especially  during  wet  weather. 
As  Mr.  Wallace  remarks,  the  natives  in  all  countries  are  glad  to 
protect  their  naked  backs  and  shoulders  with  some  slight  cover- 
ing. No  one  supposes  that  the  nakedness  of  the  skin  is  any  direct 
advantage  to  man;  his  body  therefore  cannot  have  been  divested 
of  hair  through  natural  selection.^^    Nor,  as  shown  in  a  former 

21  'ContributDons  to  the  Theory  of  Natural  Selection,'  1870,  p.  346.  Mr. 
Wallace  believes  (p.  350)  "that  some  intellig-ent  power  has  guided  or 
"determined  the  development  of  man;"  and  he  considers  the  hairless 
condition  of  the  skin  as  coming-  under  this  head.  The  Rev.  T.  R.  Steb- 
bing,  in  commenting  on  this  view  ('Transactions  of  Devonshire  Assoc, 
for  Science,'  1870)  remarks,  that  had  Mr.  Wallace  "employed  his  usual 
"ingenuity  on  the  question  of  man's  hairless  skin,  he  might  have  seen 


MAN— ABSENCE  OF  HAIR.  595 

chapter,  have  we  any  evidence  that  this  can  be  due  to  the 
direct  action  oi  climate,  or  that  it  is  the  result  of  correlated  de- 
velopment. 

The  absence  of  hair  on  the  body  is  to  a  certain  extent  a  sec- 
ondary sexual  character;  for  in  all  parts  of  the  world  women 
are  less  hairy  than  men.  Therefore  we  may  reasonably  suspect 
that  this  character  has  been  gained  through  sexual  selection. 
We  know  that  the  faces  of  several  species  of  monkeys,  and  large 
surfaces  at  the  posterior  end  of  the  body  of  other  species,  have 
been  denuded  of  hair;  and  this  we  may  safely  attribute  to  sexual 
selection,  for  these  surfaces  are  not  only  vividly  colored,  but  some- 
times, as  with  the  male  mandrill  and  female  rhesus,  much  more 
vividly  in  the  one  sex  than  in  the  other,  especially  during  the 
breeding-season.  I  am  informed  by  Mr.  Bartlett  that,  as  these 
animals  gradually  reach  maturity,  the  naked  surfaces  grow  larger 
compared  with  the  size  of  their  bodies.  The  hair,  however,  ap- 
pears to  have  been  removed,  not  for  the  sake  of  nudity,  but  that 
the  color  of  the  skin  may  be  more  fully  displayed.  So  again  with 
many  birds,  it  appears  as  if  the  head  and  neck  had  been  divested 
of  feathers  through  sexual  selection,  to  exhibit  the  brightly-col- 
ored skin. 

As  the  body  in  woman  is  less  hairy  than  in  man,  and  as  this 
character  is  common  to  all  races,  we  may  conclude  that  it  was 
our  female  semi-human  ancestors  who  were  first  divested  of  hair, 
and  that  this  occurred  at  an  extremely  remote  period  before  the 
several  races  had  diverged  from  a  common  stock.  Whilst  our 
female  ancestors  were  gradually  acquiring  this  new  character  of 
nudity,  they  must  have  transmitted  it  almost  equally  to  their 
offspring  of  both  sexes  whilst  young;  so  that  its  transmission, 
as  with  the  ornaments  of  many  mammals  and  birds,  has  not  been 
limited  either  by  sex  or  age.  There  is  nothing  surprising  in  a 
partial  loss  of  hair  having  been  esteemed  as  an  ornament  by  our 
ape-like  progenitors,  for  we  have  seen  that  innumerable  strange 
characters  have  been  thus  esteemed  by  animals  of  all  kinds,  and 
have  consequently  been  gained  through  sexual  selection.  Nor 
is  it  surprising  that  a  slightly  injurious  character  should  have 
been  thus  acquired;  for  we  know  that  this  is  the  case  with  the 
plumes  of  certain  birds,  and  with  the  horns  of  certain  stags. 

The  females  of  some  of  the  anthropoid  apes,  as  stated  in  a 
former  chapter,  are  somewhat  less  hairy  on  the  under  surface  than 
the  males;  and  here  we  have  what  might  have  afforded  a  com- 
mencement for  the  process  of  denudation.  With  respect  to  the 
completion  of  the  process  through  sexual  selection,  it  is  well  to 
bear  in  mind  the  New  Zealand  proverb,  "There  is  no  woman  for 

"the  possibility   of   its    selection   through  its   superior   beauty   or  the 
"health  attaching  to  superior  cleanliness." 


&96  THE  DESCENT  OF  MAN. 

"a  hairy  man."  All  who  have  seen  photographs  of  the  Siamese 
hairy  family  will  admi^,  how  ludicrously  hideous  is  the  opposite 
extreme  of  excessive  hairiness.  And  the  king  of  Siam  had  to 
bribe  a  man  to  marry  the  first  hairy  woman  in  the  family;  ana 
she  transmitted  this  character  to  her  young  offspring  of  both 
sexes.^ 

Some  races  are  much  more  hairy  than  others,  especially  the 
males;  but  it  must  not  be  assumed  that  the  more  hairy  races, 
such  as  the  European,  have  retained  their  primordial  condition 
more  completely  than  the  naked  races,  such  as  the  Kalmucks  or 
Americans.  It  is  more  probable  that  the  hairiness  of  the  former 
is  due  to  partial  reversion;  for  characters  which  have  been  at 
some  former  period  long  inheritea,  are  always  apt  to  return.  We 
have  seen  that  idiots  are  often  very  hairy,  and  they  are  apt  to 
revert  in  other  characters  to  a  lower  animal  type.  It  does  not 
appear  that  a  cold  cliniate  has  been  influential  in  leading  to  this 
kind  of  reversion;  excepting  perhaps  with  the  negroes,  who  have 
been  reared  during  several  generations  in  the  United  States,-^ 
and  possibly  with  the  Ainos,  who  inhabit  the  northern  islands  of 
the  Japan  Archipelago.  But  the  laws  of  inheritance  are  so  com- 
plex that  we  can  seldom  understand  their  action.  If  the  greater 
hairiness  of  certain  races  be  the  result  of  reversion,  unchecked  by 
any  form  of  selection,  its  extreme  variability,  even  within  the 
limits  of  the  same  race,  ceases  to  be  remarkable. 

With  respect  to  the  bearu  in  man,  if  we  turn  to  our  best  guide, 
the  Quadrumana,  we  find  beards  equally  developed  in  both  sex8S 

22  'The  Variation  of  Animals  and  Plants  under  Domestication,'  vol.  ii. 
1868,   p.   327. 

23  'Investigations  into  Military  and  Anthropological  Statistics  of 
American  Soldiers,'  by  B.  A.  Gould,  1869;  p.  568:— Observations  were 
carefully  made  on  the  ^xairiness  (  2129  black  and  colored  soldiers,  whilst 
they  were  bat.iing;  and  by  looking  to  the  published  table,  "it  is  mani- 
"fest  at  a  glance  that  there  is  but  little,  if  any,  difference  between  the 
"white  and  the  black  races  in  this  respect."  It  is,  however,  certain, 
that  negroes  in  their  native  and  much  hotter  land  of  Africa,  have 
remarkably  smooth  bodies.  It  should  be  particularly  observed,  that 
both  pure  blacks  and  mulattoes  were  included  in  the  above  enumer- 
ation; and  this  is  an  unfortunate  circumstance,  as  in  accordance  with 
a  principle,  the  truth  of  which  I  have  elsewhere  proved,  crossed  races 
of  man  would  be  eminently  liable  to  revert  to  the  primordial  hairy 
character  of  their  early  ape-like  progenitors. 

2*  Hardly  any  view  advanced  in  this  work  has  met  with  so  much 
disfavor  (see  for  instance,  Spengel,  'Die  Portschritte  des  Darwinismus,' 
1874,  p.  80)  as  the  above  explanation  of  the  loss  of  hair  in  mankind 
through  sexual  selection;  but  none  of  the  opposed  arguments  seem  to 
me  of  much  weight,  in  comparison  with  the  facts  showing  that  the 
nudity  of  the  skin  is  to  a  certain  extent  a  secondary  sexual  character 
in  naan  and  in  some  of  the  Quadrumana. 


BEARDS.  597 

of  many  species,  but  in  some,  either  confined  to  the  males,  or 
more  developed  in  them  than  in  the  females.  From  this  fact  and 
from  the  curious  arrangement,  as  well  as  the  bright  colors  of  the 
hair  about  the  heads  of  many  monkeys,  it  is  highly  probable,  as 
before  explained,  that  the  males  first  acquired  their  beards 
through  sexual  selection  as  an  ornament,  transmitting  them  in 
most  cases,  equally  or  nearly  so,  to  their  offspring  of  both  sexes. 
We  know  from  Eschricht-"  that  with  mankind,  the  female  as  well 
as  the  male  foetuo  is  furnished  with  much  hair  on  the  face,  espe- 
cially round  the  mouth;  and  this  indicates  that  we  are  descended 
from  progenitors,  of  whom  both  sexes  were  bearded.  It  appears 
therefore  at  first  sight  probable  than  man  has  retained  his  beard 
from  a  very  early  period,  whilst  woman  lost  her  beard  at  the 
same  time  that  her  body  became  almost  completely  divested  of 
hair.  Even  the  color  of  our  beards  seems  to  have  been  inherited 
from  an  ape-like  progenitor;  for  when  there  is  any  difference  in 
tint  between  the  hair  of  the  head  and  the  beard,  the  latter  is 
lighter  colored  in  all  monkeys  and  in  man.  In  those  Quadrumana 
in  which  the  male  has  a  larger  beard  than  that  of  the  female,  it 
is  fully  developed  only  at  maturity,  just  as  with  mankind;  and 
it  is  possible  that  only  the  later  stages  of  development  have  been 
retained  by  man.  In  opposition  to  this  view  of  the  retention  cf 
the  beard  from  an  early  period,  is  the  fact  of  its  great  variability 
in  different  races,  and  even  within  the  same  race;  for  this  indi- 
cates reversion, — long  lost  characters  being  very  apt  to  vary  on 
reappearance. 

Nor  must  we  overlook  the  part  which  sexual  selection  may 
have  played  in  later  times;  for  we  know  that  with  savages,  the 
men  of  the  beardless  races  take  infinite  pains  in  eradicating  every 
hair  from  their  faces  as  something  odious,  whilst  the  men  of 
the  bearded  races  feel  the  greatest  pride  in  their  beards.  The 
women,  no  doubt,  participate  in  these  feelings,  and  if  so  sexual 
selection  can  hardly  have  failed  to  have  effected  something  in  the 
course  of  later  times.  It  is  also  possible  that  the  long-continued 
habit  of  eradicating  the  hair  may  have  produced  an  inherited 
effect.  Dr.  Brown-Sequard  has  shown  that  if  certain  animals  are 
operated  on  in  a  particular  manner,  their  offspring  are  affected. 
Further  evidence  could  be  given  of  the  inheritance  of  the  effects 
of  mutilations;  but  a  fact  lately  ascertained  by  Mr.  Salvin^^  has 
a  more  direct  bearing  on  the  present  question;  for  he  has  shown 
that  the  motmots,  which  are  known  habitually  to  bite  off  the  barbs 
of  the  two  central  tail-feathers,  have  the  barbs  of  these  feathers 


25  'Ueber  die  Richtung  der  Haare  am  Menschlichen  Korper,'  in  Mul- 
ler's  'Archiv  fur  Anat.  und  Phys.'  1837,  s.  40. 

28  'On  the  tail-feathers  of  Momotus'  'Proc.  Zoolog.  Soc.,'  1873,  p.  429. 

39 


598  -THE  DESCENT  OP  MAN. 

naturally  somewhat  reduced.-^  Nevertheless  with  mankind,  the 
habit  of  eradicating  the  beard  and  the  hairs  on  the  body  would 
probably  not  have  arisen  until  these  had  already  become  by  some 
means  reduced. 

It  is  difficult  to  form  any  judgment  as  to  how  the  hair  on  the 
head  became  developed  to  its  present  great  length  in  many  races. 
Eschricht-®  states  that  in  the  human  fcstus  the  hair  on  the  face 
during  the  fifth  month  is  longer  than  that  on  the  head;  and 
this  indicates  that  our  semi-human  progenitors  were  not  fur- 
nished v/ith  long  tresses,  which  must  therefore  have  been  a  late 
acquisition.  This  is  likewise  indicated  by  the  extraordinary  dif- 
ference in  the  length  of  the  hair  in  the  different  races;  in  the 
negro  the  hair  forms  a  mere  curly  mat;  v/ith  us  it  is  of  great 
length,  and  with  the  American  natives  it  not  rarely  reaches  to 
the  ground.  Some  species  of  Semnopithecus  have  their  heads 
covered  with  moderately  long  hair,  and  this  probably  serves  as 
an  ornament  and  was  acquired  through  sexual  selection.  The 
same  view  may  perhaps  be  extended  to  mankind,  for  vre  knov/ 
that  long  tresses  are  now  and  were  formerly  much  admired,  as 
may  be  observed  in  the  works  of  almost  every  poet;  St.  Paul 
says,  "if  a  woman  have  long  hair,  it  is  a  glory  to  her;"  and  we 
have  seen  that  in  North  America  a  chief  was  elected  solely  from 
the  length  of  his  hair. 

Color  of  the  Skin.  — The  best  kind  of  evidence  that  in  man  the 
color  of  the  skin  has  been  modified  through  sexual  selection  is 
scanty;  for  in  most  races  the  sexes  do  not  differ  in  this  respect, 
and  only  slightly,  as  we  have  seen,  in  others.  We  know,  however, 
from  the  many  facts  already  given  that  the  color  of  the  skin  is 
regarded  by  the  men  of  all  races  as  a  highly  important  element 
in  their  beauty;  so  that  it  is  a  character  which  would  be  likely 
to  have  been  modified  through  selection,  as  has  occurred  in  in- 
numerable instances  with  the  lower  animals.  It  seems  at  first 
sight  a  monstrous  supposition  that  the  jet-blackness  of  the  negro 
should  have  been  gained  through  sexual  selection;  but  this  view 
is  supported  by  various  analogies,  and  we  know  that  negroes  ad- 
mire their  ov^rn  color.  With  mammals,  v/hen  the  sexes  differ  in 
color,  the  male  is  often  black  or  much  darker  than  the  female; 
and  it  depends  merely  on  the  form  of  inheritance  whether  this  or 
any  other  tint  is  transmitted  to  both  sexes  or  to  one  alone.  The 
resemblance  to  a  negro  in  miniature  of  Pithecia  satanas  with  his 


27  Mr.  Sproat  has  suggested  ('Scenes  and  Studies  of  Savage  Life,' 
1868,  p.  25)  this  same  view.  Some  distinguished  ethnologists,  amongst 
others  M.  Gosse  of  Geneva,  believe  that  artificial  modifications  of  the 
skull  tend  to  be  inherited. 

28  'Ueber  die  Richtung,'  ibid.  s.  49. 


SUMMARY  ON  MAN. 

jet  black  skin,  white  rolling  eyeballs,  and  hair  parted  on  the  top 
of  the  head,  is  almost  ludicrous. 

The  color  of  the  face  differs  much  more  widely  in  the  various 
kinds  of  monkeys  than  it  does  in  the  races  of  man;  and  we  have 
some  reason  to  believe  that  the  red,  blue,  orange,  almost  white 
and  black  tints  of  their  skin,  even  when  common  to  both  sexes, 
as  well  as  the  bright  colors  of  their  fur,  and  the  ornamental 
tufts  about  the  head,  have  all  been  acquired  through  sexual  selec- 
tion. As  the  order  of  development  during  growth,  generally  in- 
dicates the  order  in  which  the  characters  of  a  species  have  been 
developed  and  modified  during  previous  generations;  and  as  the 
newly-born  infants  of  the  various  races  of  man  do  not  differ 
nearly  as  much  in  color  as  do  the  adults,  although  their  bodies 
are  as  completely  destitute  of  hair,  we  have  some  slight  evidence 
that  the  tints  of  the  different  races  were  acquired  at  a  period  sub- 
sequent to  the  removal  of  the  hair,  which  must  have  occurred  at 
a  very  early  period  in  the  history  of  man. 

Suwiinavy. — We  may  conclude  that  the  greater  size,  strength, 
courage,  pugnacity,  and  energy  of  man,  in  comparison  with  wo- 
man, were  acquired  during  primeval  times,  and  have  subsequently 
been  augmented,  chiefly  through  the  contests  of  rival  males  for 
the  possession  of  the  females.  The  greater  intellectual  vigor  and 
power  of  invention  in  man  is  probably  due  to  natural  selection, 
combined  with  the  inherited  effects  of  habit,  for  the  most  able 
men  will  have  succeeded  best  in  defending  and  providing  for  them- 
selves and  for  their  wives  and  offspring.  As  far  as  the  extreme 
intricacy  of  the  subject  permits  us  to  judge,  it  appears  that  our 
male  ape-like  progenitors  acquired  their  beards  as  an  ornament 
to  charm  or  excite  the  opposite  sex,  and  transmitted  them  only 
to  their  male  offspring.  The  females  apparently  first  had  their 
bodies  denuded  of  hair,  also  as  a  sexual  ornament;  but  they  trans- 
mitted this  character  almost  equally  to  both  sexes.  It  is  not  im- 
probable that  the  females  were  modified  in  other  respects  for  the 
same  purpose  and  by  the  same  means;  so  that  women  have  ac- 
quired sweeter  voices  and  become  more  beautiful  than  men. 

It  deserves  attention  that  with  mankind  the  conditions  were  in 
many  respects  much  more  favorable  for  sexual  selection,  during 
a  very  early  period,  when  man  had  only  just  attained  to  the  rank 
of  manhood,  than  during  later  times.  For  he  would  then,  as  we 
may  safely  conclude,  have  been  guided  more  by  his  instinctive 
passions,  and  less  by  foresight  or  reason.  He  would  have  jealously 
guarded  his  wife  or  wives.  He  would  not  have  practiced  infanti- 
cide; nor  valued  his  wives  merely  as  useful  slaves;  nor  have  been 
betrothed  to  them  during  infancy.  Hence  we  may  infer  that  the 
races  of  men  were  differentiated  as  far  as  sexual  selection  is  con- 
cerned, in  chief  part  at  a  very  remote  epoch;    and  this  conclusion 


600  THE  DESCENT  OF  MAN. 

throws  light  on  the  remarkable  fact  that  at  the  most  ancient  peri- 
od, of  which  we  have  as  yet  any  record,  the  races  of  man  had 
already  come  to  differ  nearly  or  quite  as  much  as  they  do  at  the 
present  day. 

The  views  here  advanced,  on  the  part  which  sexual  selection 
has  played  in  the  history  of  man,  want  scientific  precision.  He 
who  does  not  admit  this  agency  in  the  case  of  the  lower  animals, 
will  disregard  all  that  I  have  written  in  the  later  chapters  on 
man.  We  cannot  positively  say  that  this  character,  but  not  that, 
has  been  thus  modified;  it  has,  however,  been  shown  that  the 
races  of  man  differ  from  each  other  and  from  their  nearest  allies, 
in  certain  characters  which  are  of  no  service  to  them  in  their  daily 
habits  of  life,  and  which  it  is  extremely  probable  would  have  been 
modified  through  sexual  selection.  We  have  seen  that  with  the 
lowest  savages  the  people  of  each  tribe  admire  their  own  charac- 
teristic qualities, — the  shape  of  the  head  and  face,  the  square- 
ness of  the  cheek-bones,  the  prominence  or  depression  of  the  nose, 
the  color  of  the  skin,  the  length  of  the  hair  on  the  head,  the  ab- 
sence of  hair  on  the  face  and  body,  or  the  presence  of  a  great 
beard,  and  so  forth.  Hence  these  and  other  such  points  could 
hardly  fail  to  be  slowly  and  gradually  exaggerated,  from  the  more 
powerful  and  able  men  in  each  tribe,  who  would  succeed  in  rear- 
ing the  largest  number  of  offspring,  having  selected  during  many 
generations  for  their  wives  the  most  strongly  characterized  and 
therefore  most  attractive  women.  For  my  own  part  I  conclude 
that  of  all  the  causes  which  have  led  to  the  differences  in  external 
appearance  between  the  races  of  man,  and  to  a  certain  extent 
between  man  and  the  lower  animals,  sexual  selection  has  been 
the  most  efficient. 


GENERAL  SUMMARY.  «01 


CHAPTER    XXI. 

GENERAL  SUMMARY  AND  CONCLUSION. 

Main  conclusion  that  man  is  descended  from  some  lower  form— Manner 
of  development— Genealogy  of  man— Intellectual  and  moral  facul- 
ties— Sexual  selection — Concluding-  remarks. 

A  brief  summary  will  be  sufficient  to  recall  to  the  reader's  mind 
the  more  salient  points  in  this  work.  Many  of  the  views  which 
have  been  advanced  are  highly  speculative,  and  some  no  doubt 
will  prove  erroneous;  but  I  have  in  every  case  given  the  reasons 
which  have  led  me  to  one  view  rather  than  to  another.  It 
seemed  worth  while  to  try  how  far  the  principle  of  evolution 
would  throw  light  on  some  of  the  more  complex  problems  in  the 
natural  history  of  man.  False  facts  are  highly  injurious  to  the 
progress  of  science,  for  they  often  endure  long;  but  false  views, 
if  supported  by  some  evidence,  do  little  harm,  for  every  one  takes 
a  salutary  pleasure  in  proving  their  falseness;  and  when  this  is 
done,  one  path  towards  error  is  closed  and  the  road  to  truth  is 
often  at  the  same  time  opened. 

The  main  conclusion  here  arrived  at,  and  now  held  by  many 
naturalists  who  are  well  competent  to  form  a  sound  judgment,  is 
that  man  is  descended  from  some  less  highly  organized  form.  The 
grounds  upon  which  this  conclusion  rests  will  never  be  shaken, 
for  the  close  similarity  between  man  and  the  lower  animals  in 
embryonic  development,  as  well  as  in  innumerable  points  of  struc- 
ture and  constitution,  both  of  high  and  of  the  most  trifling  im- 
portance,— the  rudiments  which  he  retains,  and  the  abnormal 
reversions  to  which  he  is  occasionally  liable, — are  facts  which  can- 
not be  disputed.  They  have  long  been  known,  but  until  recently 
they  told  us  nothing  with  respect  to  the  origin  of  man.  Now 
when  viewed  by  the  light  of  our  knowledge  of  the  whole  organic 
world,  their  meaning  is  unmistakable.  The  great  principle  of 
evolution  stands  up  clear  and  firm,  when  these  groups  of  facts 
are  considered  in  connection  with  others,  such  as  the  mutual  af- 
finities of  the  members  of  the  same  group,  their  geographical  dis- 
tribution in  past  and  present  times,  and  their  geological  succes- 
sion.   It  is  incredible  that  all  these  facts  should  speak  falsely. 


602  THE   DESCENT  OF  MAN. 

He  who  is  not  content  to  look,  like  a  savage,  at  the  phenomena 
of  nature  as  disconnected,  cannot  any  longer  believe  that  man  is 
the  work  of  a  separate  act  of  creation.  He  will  be  forced  to  admit 
that  the  close  resemblance  of  the  embryo  of  man  to  that,  for  in- 
stance of  a  dog — the  construction  of  his  skull,  limbs  and  whole 
frame  on  the  same  plan  with  that  of  other  mammals,  independent- 
ly of  the  uses  to  which  the  parts  may  be  put — the  occasional  re- 
appearance of  various  structures,  for  instance  of  several  muscles^ 
which  man  does  not  normally  possess,  but  which  are  common  to 
the  Quadrumana — and  a  crowd  of  analogous  facts— all  point  in 
the  plainest  manner  to  the  conclusion  that  man  is  the  co-descend- 
ant with  other  mammals  of  a  common  progenitor. 

We  have  seen  that  man  incessantly  presents  individual  differ- 
ences in  all  parts  of  his  body  and  in  his  mental  faculties.  These 
differences  or  variations  seem  to  be  induced  by  the  same  general 
causes,  and  to  obey  the  same  laws  as  with  the  lower  animals. 
In  both  cases  similar  lav/s  of  inheritance  prevail.  Man  tends  to 
increase  at  a  greater  rate  than  his  means  of  subsistence;  con- 
sequently he  is  occasionally  subjected  to  a  severe  struggle  for 
existence,  and  natural  selection  will  have  effected  whatever  lies 
within  its  scope.  A  succession  of  strongly-marked  variations  of  a 
similar  nature  is  by  no  means  requisite;  slight  fluctuating  differ- 
ences in  the  individual  suffice  for  the  work  of  natural  selection; 
not  that  we  have  any  reason  to  suppose  that  in  the  same  species, 
all  parts  of  the  organization  tend  to  vary  to  the  same  degree. 
We  may  feel  assured  that  the  inherited  effects  of  the  long-con- 
tinued use  or  disuse  of  parts  will  have  done  much  in  the  same 
direction  with  natural  selection.  Modifications  formerly  of  im- 
portance, though  no  longer  of  any  special  use,  are  long-inherited. 
When  one  part  is  modified,  other  parts  change  through  the  prin- 
ciple of  correlation,  of  which  we  have  instances  in  many  curious 
cases  of  correlated  monstrosities.  Something  may  be  attributed 
to  the  direct  and  definite  action  of  the  surrounding  conditions  of 
life,  such  as  abundant  food,  heat  or  moisture;  and  lastlj^,  many 
characters  of  slight  physiological  importance,  some  indeed  of  con- 
siderable importance,  have  been  gained  through  sexual  selection. 

No  doubt  man,  as  well  as  every  other  animal,  presents  struct- 
ures, which  seem  to  our  limited  knowledge,  not  to  be  now  of  any 
service  to  him,  nor  to  have  been  so  formerly,  either  for  the  gen- 
eral conditions  of  life,  or  in  the  relations  of  one  sex  to  the  other. 
Such  structures  cannot  be  accounted  for  by  any  form  of  selection, 
or  by  the  inherited  effects  of  the  use  and  disuse  of  parts.  W^e 
know,  however,  that  many  strange  and  strongly-marked  peculiar- 
ities of  structure  occasionally  appear  in  our  domesticated  produc- 
tions, and  if  their  unknov/n  causes  v»rere  to  act  more  uniformly, 
they  would  probably  become  common  to  ail  the  individuals  of 
the  species.    We  may  hope  hereafter  to  understand  something 


GENERAL.  SUMMARY.  603 

about  the  causes  of  such  occasional  modifications,  especially 
through  the  study  of  monstrosities:  hence  the  labors  of  experi- 
mentalists, such  as  those  of  M.  Camille  Dareste,  are  full  of 
promise  for  the  future.  In  general  we  can  only  say  that  the 
cause  of  each  slight  variation  and  of  each  monstrosity  lies  much 
more  in  the  constitution  of  the  organism,  than  in  the  nature  of 
the  surrounding  conditions;  though  new  and  changed  conditions 
certainly  play  an  important  part  in  exciting  organic  changes  of 
many  kinds. 

Through  the  means  just  specified,  aided  perhaps  by  others  as 
yet  undiscovered,  man  has  been  raised  to  his  present  state. 
But  since  he  attained  to  the  rank  of  manhood,  he  has  diverged 
into  distinct  races,  or  as  they  may  be  more  fitly  called,  sub- 
species. Some  of  these,  such  as  the  Negro  and  European,  are 
so  distinct  that,  if  specimens  had  been  brought  to  a  naturalist 
without  any  further  information,  they  would  undoubtedly  have 
been  considered  by  him  as  good  and  true  species.  Nevertheless 
all  the  races  agree  in  so  many  unimportant  details  of  structure 
and  in  so  many  mental  peculiarities,  that  these  can  be  accounted 
for  only  by  inheritance  from  a  common  progenitor;  and  a  pro- 
genitor thus  characterized  would  probably  deserve  to  rank  as  man. 

It  must  not  be  supposed  that  the  divergence  of  each  race  from 
the  other  races,  and  of  all  from  a  common  stock,  can  be  traced 
back  to  any  one  pair  of  progenitors.  On  the  contrary,  at  every 
stage  in  the  process  of  modification,  all  the  individuals  which 
were  in  any  way  better  fitted  for  their  conditions  of  life,  though 
in  different  degrees,  would  have  survived  in  greater  numbers 
than  the  less  well-fitted.  The  process  would  have  been  like  that 
followed  by  man,  when  he  does  not  intentionally  select  particular 
individuals,  but  breeds  from  all  the  superior  individuals,  and 
neglects  the  inferior.  He  thus  slowly  but  surely  modifies  his 
stock,  and  unconsciously  forms  a  new  strain.  So  with  respect 
to  modifications  acquired  independently  of  selection,  and  due 
to  variations  arising  from  the  nature  of  the  organism  and  the 
action  of  the  surrounding  conditions,  or  from  changed  habits  ci 
life,  no  single  pair  will  have  been  modified  much  more  than  the 
other  pairs  inhabiting  the  same  country,  for  all  will  have  been 
continually  blended  through  free  intercrossing. 

By  considering  the  embryological  structure  of  man, — the  homo- 
logies which  he  presents  with  the  lower  animals, — the  rudi- 
ments which  he  retains, — and  the  reversions  to  which  he  is  liable, 
we  can  partly  recall  in  imagination  the  former  condition  of  our 
early  progenitors;  and  can  approximately  place  them  in  their 
proper  place  in  the  zoological  series.  We  thus  learn  that  man  ig 
descended  from  a  hairy,  tailed  quadruped,  probably  arboreal  in 
its  habits,  and  an  inhabitant  of  the  Old  World.  This  creature, 
if  its  whole  structure  had  been  examined  by  a  naturalist,  would 


604  THE  DESCENT  OF  MAN. 

have  been  classed  amongst  the  Quadrumana,  as  surely  as  th® 
still  more  ancient  progenitor  of  the  Old  and  New  World  monkeys. 
The  Quadrumana  and  all  the  higher  mammals  are  probably  de- 
rived from  an  ancient  marsupial  animal,  and  this  through  a 
long  line  of  diversified  forms,  from  some  amphibian-like  creature, 
and  this  again  from  som^^  fish-like  animal.  In  the  dim  obscurity 
of  the  past  we  can  see  that  the  early  progenitor  of  all  the  Verte* 
brata  must  have  been  an  aquatic  animal,  provided  with  branchiae, 
with  the  two  sexes  united  in  the  same  individual,  and  with  the 
most  important  organs  of  the  body  (such  as  the  brain  and  heart) 
imperfectly  or  not  at  all  developed.  This  animal  seems  to  have 
been  more  like  the  larvae  of  the  existing  marine  Ascidians  than 
any  other  known  form. 

The  high  standard  ot  our  intellectual  powers  and  moral  dis- 
position is  the  greatest  difficulty  which  presents  itself,  after  we 
have  been  driven  to  this  conclusion  on  the  origin  of  man.  (But 
every  one  who  admits  the  principle  of  evolution,  must  see  that 
the  mental  powers  of  tlia  higher  animals,  which  are  the  same  in 
kind  with  those  of  man,  though  so  different  in  degree,  are  capabl? 
of  advancement.  Thus  the  interval  between  the  mental  powers 
of  one  of  the  higher  apes  and  of  a  fish,  or  between  those  of  an 
ant  and  scale-insect,  is  immense;  yet  their  development  does  not 
offer  any  special  difficulty;  for  with  our  domesticated  animals, 
the  mental  faculties  are  certainly  variable,  and  the  variations 
are  inherited.  No  one  doubts  that  they  are  of  the  utmost  im- 
portance to  animals  in  a  state  of  nature.  Therefore  the  condi- 
tions are  favorable  for  their  development  through  natural  selec- 
tion. The  same  conclusion  may  be  extended  to  man;  the  intel- 
lect must  have  been  all-important  to  him,  even  at  a  very  remote 
period,  as  enabling  him  to  invent  and  use  language,  to  make 
weapons,  tools,  traps,  &c.,  whereby  with  the  aid  of  his  social 
habits,  he  long  ago  became  the  most  dominant  of  all  living  crea- 
tures^ji 

A  great  stride  in  the  development  of  the  intellect  will  have 
followed,  as  soon  as  the  half-art  and  half-instinct  of  language 
came  into  use;  for  the  continued  use  of  language  will  have 
reacted  on  the  brain  and  produced  an  inherited  effect;  and  this 
again  will  have  reacted  on  the  improvement  of  language.  As  Mr. 
Chauncey  Wright^  has  well  remarked,  the  largeness  of  the  brain 
in  man  relatively  to  his  body,  compared  with  the  lower  animals, 
may  be  attributed  in  chief  part  to  the  early  use  of  some  simple 
form  of  language, — that  wonderful  engine  which  aflaxes  signs  to 
all  sorts  of  objects  and  qualities,  and  excites  trains  of  thought 


1  'On  the  Limits  of  Natural  Selection,'  in  the  'North  American  Re- 
'view,'  Oct.  1870,  p.  295. 


GENERAL  SUMMARY.  605 

which  would  never  arise  from  the  mere  impression  of  the  senses, 
or  if  they  did  arise,  could  not  be  followed  out.  The  higher  in- 
tellectual powers  of  man,  such  as  those  of  ratiocination,  abstrac- 
tion, self-consciousness,  &c.,  probably  follow  from  the  continued 
improvement  and  exercise  of  the  other  mental  faculties.  ^ 

The  development  of  the  moral  qualities  is  a  more  interesting! 
problem.  The  foundation  lies  in  the  social  instincts,  including 
under  this  term  the  family  ties.  These  instincts  are  highly  com- 
plex, and  in  the  case  of  the  lower  animals  give  special  tendencies 
toward  certain  definite  actions;  but  the  more  important  ele- 
ments are  love,  and  the  distinct  emotion  of  sympathy.  Animals 
endowed  with  the  social  instincts  take  pleasure  in  one  another's 
company,  warn  one  another  of  danger,  defend  and  aid  one  an- 
other in  many  ways.  These  instincts  do  not  extend  to  all  the 
individuals  of  the  species,  but  only  to  those  of  the  same  commu- 
nity. As  they  are  highly  beneficial  to  the  species,  they  have  in 
all  probability  been  acquired  through  natural  selection.  ^ 

A  moral  being  is  one  who  is  capable  of  refiecting  on  his  past 
actions  and  their  motives — of  approving  of  some  and  disapprov- 
ing of  others;  and  the  fact  that  man  is  the  one  being  who  cer- 
tainly deserves  this  designation,  is  the  greatest  of  all  distinctions 
between  him  and  the  lower  animals.  But  in  the  fourth  chapter 
I  have  endeavored  to  show  that  the  moral  sense  follows,  firstly, 
from  the  enduring  and  ever-present  nature  of  the  social  instincts; 
secondly,  from  man's  appreciation  of  the  approbation  and  dis- 
approbation of  his  fellows;  and  thirdly,  from  the  high  activity 
of  his  mental  faculties,  with  past  impressions  extremely  vivid; 
and  in  these  latter  respects  he  differs  from  the  lower  animals. 
Owing  to  this  condition  of  mind,  man  cannot  avoid  looking  both 
backwards  and  forwards,  and  comparing  past  impressions.  Hense 
after  some  temporary  desire  or  passion  has  mastered  his  social 
instincts,  he  reflects  and  compares  the  now  weakened  impression 
of  such  past  impulses  with  the  ever-present  social  instincts;  and 
he  then  feels  that  sense  of  dissatisfaction  which  all  unsatisfied  in- 
stincts leave  behind  them,  he  therefore  resolves  to  act  differently 
for  the  future, — and  this  is  conscience.  Any  instinct,  permanent- 
ly stronger  or  more  enduring  than  another,  gives  rise  to  a  feeling 
which  we  express  by  saying  that  it  ought  to  be  obeyed.  A  pointer 
dog,  if  able  to  reflect  on  his  past  conduct,  would  say  to  himself, 
I  ought  (as  indeed  we  say  of  him)  to  have  pointed  at  that  hare 
and  not  have  yielded  to  the  passing  temptation  of  hunting  it. 

Social  animals  are  impelled  partly  by  a  wish  to  aid  the  members') 
of  their  community  in  a  general  manner,  but  more  commonly  to  ' 
perform  certain  deflnite  actions.     Man  is  impelled  by  the  same 
general  wish  to  aid  his  fellows;    but  has  few  or  no  special  in- 
stincts.   He  differs  also  from  the  lower  animals  in  the  power  of 
expressing  his  desires  by  words,  which  thus  become  a  guide  to 


606  THE   DESCENT  OF  MAN. 

the  aid  required  and  bestowed.  The  motive  to  give  aid  is  likewise 
much  modified  in  man:  it  no  longer  consists  solely  of  a  blind 
instinctive  impulse,  but  is  much  influenced  by  the  praise  or  blame 
of  his  fellows.  The  appreciation  and  the  bestowal  of  praise  and 
blame  both  rest  on  sympathy;  and  this  emotion,  as  we  have 
seen,  is  one  of  the  most  important  elements  of  the  social  in- 
stincts. Sympathy,  though  gained  as  an  instinct,  is  also  much 
strengthened  by  exercise  or  habit.  As  all  men  desire  their  own 
happiness,  praise  or  blame  is  bestowed  on  actions  and  motives, 
according  as  they  lead  to  this  end;  and  as  happiness  is  an  essen- 
tial part  of  the  general  good,  the  greatest-happiness  principle  in- 
directly serves  as  a  nearly  safe  standard  of  right  and  wrong. 
As  the  reasoning  powers  advance  and  experience  is  gained,  the 
remoter  effects  of  certain  lines  of  conduct  on  the  character  of 
the  individual,  and  on  the  general  good,  are  perceived;  and  then 
the  self-regarding  virtues  come  vathin  the  scope  of  public  opin- 
ion, and  receive  praise,  and  their  opposites  blame.  But  with  the 
less  civilized  nations  reason  often  errs,  and  many  bad  customs 
and  base  superstitions  come  within  the  same  scope,  and  are  then 
esteemed  as  high  virtues,  and  their  breach  as  heavy  crimes. 

The  moral  faculties  are  generally  and  justly  esteemed  as  of 
higher  value  than  the  intellectual  powers.  But  we  should  bear 
in  mind  that  the  activity  of  the  mind  in  vividly  recalling  past 
impressions  is  one  of  the  fundamental  though  secondary  bases 
of  conscience.  This  affords  the  strongest  argument  for  educating 
and  stimulating  in  all  possible  ways  the  intellectual  faculties  of 
every  human  being.  No  doubt  a  man  with  a  torpid  mind,  if  his 
social  affections  and  sympathies  are  well  developed,  will  be  led 
to  good  actions,  and  may  have  a  fairly  sensitive  conscience.  But 
whatever  renders  the  imagination  more  vivid  and  strengthens  the 
habit  of  recalling  and  comparing  past  impressions,  will  make 
the  conscience  more  sensitive,  and  may  even  somewhat  com- 
pensate for  weak  social  affections  and  sympathies. 

The  moral  nature  of  man  has  reached  its  present  standard, 
partly  through  the  advancement  of  his  reasoning  powers  and 
conse(^uently  of  a  just  public  opinion,  but  especially  fropi  his 
sympathies  having  been  rendered  more  tender  and  widely  dif- 
fused through  the  effects  of  habit,  example,  instruction,  and  re- 
flection. It  is  not  improbable  that  after  long  practice  virtuous 
tendencies  may  be  inherited.  With  the  more  civilized  races,  the 
conviction  of  the  existence  of  an  all-seeing  Deity  has  had  a  potent 
influence  on  the  advance  of  morality.  Ultimately  man  does  not 
accept  the  praise  or  blame  of  his  fellows  as  his  sole  guide,  though 
few  escape  this  influence,  but  his  habitual  convictions,  controlled 
by  reason,  afford  him  the  safest  rule.  His  conscience  then  be- 
comes the  supreme  judge  and  monitor.  Nevertheless  the  first 
foundation  or  origin  of  the  moral  sense  lies  in  the  social  instincts^ 


CENERAL  SUMMARY.  60t 

including  sympathy;  and  these  instincts  no  doubt  were  primarily- 
gained,  as  in  the  case  of  the  lower  animals,  through  natural 
selection. 

The  belief  in  God  has  often  been  advanced  as  not  only  the"1 
greatest,  but  the  most  complete  of  all  the  distinctions  between 
man  and  the  lower  animals.  It  is  however  impossible,  as  we 
have  seen,  to  maintain  that  this  belief  is  innate  or  instinctive  in 
man.  On  the  other  hand  a  belief  in  all-pervading  spiritual  agen- 
cies seems  to  be  universal;  and  apparently  follows  from  a  con- 
siderable advance  in  man's  reason,  and  from  a  still  greater  ad- 
vance in  his  faculties  of  imagination,  curiosity  and  wonder.  I 
am  aware  that  the  assumed  instinctive  belief  in  God  has  been 
used  by  many  persons  as  an  argument  for  His  existence.  But 
this  is  a  rash  argument,  as  we  should  thus  be  compelled  to 
believe  in  the  existence  of  many  cruel  and  malignant  spirits,  only 
a  little  more  powerful  than  man;  for  the  belief  in  them  is  far 
more  general  than  in  a  beneficent  Deity.  The  idea  of  a  universal 
and  beneficent  Creator  does  not  seem  to  arise  in  the  mind  of  man, 
until  he  has  been  elevated  by  long-continued  culture. 

He  who  believes  in  the  advancement  of  man  from  some  low 
organized  form,  will  naturally  ask  how  does  this  bear  on  the  be- 
lief in  the  immortality  of  the  soul.  The  barbarous  races  of  man, 
as  Sir  J.  Lubbock  has  shown,  possess  no  clear  belief  of  this  kind; 
but  arguments  derived  from  the  primeval  beliefs  of  savages  are, 
as  we  have  just  seen,  of  little  or  no  avail.  Few  persons  feel  any 
anxiety  from  the  impossibility  of  determining  at  what  precise 
period  in  the  development  of  the  individual,  from  the  first  trace 
of  a  minute  germinal  vesicle,  man  becomes  an  immortal  being; 
and  there  is  no  greater  cause  for  anxiety  because  the  period 
cannot  possibly  be  determined  in  the  gradually  ascending  organic 
scale.2 

I  am  aware  that  the  conclusions  arrived  at  in  this  work] 
will  be  denounced  by  some  as  highly  irreligious;  but  he  who 
denounces  them  is  bound  to  show  why  it  is  more  irreligious  to 
explain  the  origin  of  man  as  a  distinct  species  by  descent  from 
some  lower  form,  through  the  laws  of  variation  and  natural 
selection,  than  to  explain  the  birth  of  the  individual  through  the 
laws  of  ordinary  reproduction.  The  birth  both  of  the  species 
and  of  the  individual  are  equally  parts  of  that  grand  sequence 
of  events,  which  our  minds  refuse  to  accept  as  the  result  of  . 
blind  chance.  The  understanding  revolts  at  such  a  conclusion, 
whether  or  not  we  are  able  to  believe  that  every  slight  variation 
of  structure, — the  union  of  each  pair  in  marriage, — the  dissem- 

-  The  Rev.  J.  A.  Picton  gives  a  discussion  to  this  effect  in  his  'New 
Theories  and  the  Old  Faith,'  1870. 


608  THE  DESCENT  OF  MAN. 

iuation  of  each  seed, — and  other  such  events,  have  all  been  or- 
dained for  some  special  purpose. 

Sexual  selection  has  been  treated  at  great  length  in  this  work; 
for,  as  I  have  attempted  to  show,  it  has  played  an  important  part 
in  the  history  of  the  organic  world.  I  am  aware  that  much  re- 
mains doubtful,  but  I  have  endeavored  to  give  a  fair  view  of  the 
whole  case.  In  the  lower  divisions  of  the  animal  kingdom,  sexual 
selection  seems  to  have  done  nothing:  such  animals  are  often 
af&xed  for  life  to  the  same  spot,  or  have  the  sexes  combined  in 
the  same  individual,  or  what  is  still  more  important,  their  per- 
ceptive and  intellectual  faculties  are  not  sufficiently  advanced  to 
allow  of  the  feelings  of  love  and  jealousy,  or  of  the  exertion  of 
choice.  When,  however,  we  come  to  the  Arthropoda  and  Verte- 
brata,  even  to  the  lowest  classes  in  these  two  great  Sub-Kingdoms, 
sexual  selection  has  effected  much. 

In  the  several  great  classes  of  the  animal  kingdom, — in  mam- 
mals, birds,  reptiles,  fishes,  insects,  and  even  crustaceans, — the  dif- 
ferences between  the  sexes  follow  nearly  the  same  rules.  The 
males  are  almost  always  the  wooers;  and  they  alone  are  armed 
with  special  weapons  for  fighting  with  their  rivals.  They  are 
generally  stronger  and  larger  than  the  females,  and  are  endowed 
with  the  requisite  qualities  of  courage  and  pugnacity.  They  are 
provided,  either  exclusively  or  in  a  much  higher  degree  than  the 
females,  with  organs  for  vocal  or  instrumental  music,  and  vrith 
odoriferous  glands.  They  are  ornamented  with  infinitely  diversi- 
fied appendages,  and  with  the  most  brilliant  or  conspicuous  col- 
ors, often  arranged  in  elegant  patterns,  whilst  the  females  are 
unadorned.  When  the  sexes  differ  in  more  important  structures, 
it  is  the  male  which  is  provided  with  special  sense-organs  for 
discovering  the  female,  with  locomotive  organs  for  reaching  her, 
and  often  with  prehensile  organs  for  holding  her.  These  various 
structures  for  charming  or  securing  the  female  are  often  devel- 
oped in  the  male  during  only  part  of  the  year,  namely  the  breed- 
ing-season. They  have  in  many  cases  been  more  or  less  trans- 
ferred to  the  females;  and  in  the  latter  case  they  often  appear 
in  her  as  mere  rudiments.  They  are  lost  or  never  gained  by  the 
males  after  emasculation.  Generally  they  are  not  developed  in 
the  male  during  early  youth,  but  appear  a  short  time  before  the 
age  for  reproduction.  Hence  in  most  cases  the  young  of  both  sexes 
resemble  each  other;  and  the  female  somewhat  resembles  her 
young  offspring  throughout  life.  In  almost  every  great  class  a 
few  anomalous  cases  occur,  where  there  has  been  an  almost  com- 
plete transposition  of  the  characters  proper  to  the  two  sexes;  the 
females  assuming  characters  which  properly  belong  to  the  males. 
This  surprising  uniformity  in  the  laws  regulating  the  differences 
between  the  sexes  in  so  many  and  such  widely  separated  classes. 


GKNERAli  SUMMARY.  609 

is  intelligible  if  we  admit  the  action  of  one  common  cause,  namely- 
sexual  selection. 

Sexual  selection  depends  on  the  success  of  certain  individuals 
over  others  of  the  same  sex,  in  relation  to  the  propagation  of  the 
species;  whilst  natural  selection  depends  on  the  success  of  both 
sexes  at  all  ages,  in  relation  to  the  general  conditions  of  life. 
The  sexual  struggle  is  of  two  kinds;  in  the  one  it  is  between  the 
individuals  of  the  same  sex,  generally  the  males,  in  order  to  drive 
away  or  kill  their  rivals,  the  females  remaining  passive;  whilst 
in  the  other,  the  struggle  is~  likewise  between  the  individuals  of 
the  same  sex,  in  order  to  excite  or  charm  those  of  the  opposite 
sex,  generally  the  females,  which  no  longer  remain  passive,  but 
select  the  more  agreeable  partners.  This  latter  kind  of  selection 
is  closely  analogous  to  that  which  man  unintentionally,  yet  ef- 
fectually, brings  to  bear  on  his  domesticated  productions,  when 
he  preserves  during  a  long  period  the  most  pleasing  or' useful 
individuals,  without  any  wish  to  modify  the  breed. 

The  laws  of  inheritance  determine  whether  characters  gained 
through  sexual  selection  by  either  sex  shall  be  transmitted  to  the 
same  sex,  or  to  both;  as  well  as  the  age  at  which  they  shall  be 
developed.  It  appears  that  variations  arising  late  in  life  are 
commonly  transmitted  to  one  and  the  same  sex.  Variability  is 
the  necessary  basis  for  the  action  of  selection,  and  is  wholly 
independent  of  it.  It  follows  from  this,  that  variations  of  the 
same  general  nature  have  often  been  taken  advantage  of  and 
accumulated  through  sexual  selection  in  relation  to  the  propaga- 
tion of  the  species,  as  well  as  through  natural  selection  in  relation 
to  the  general  purposes  of  life.  Hence  secondary  sexual  char- 
acters, when  equally  transmitted  to  both  sexes  can  be  distin- 
guished from  ordinary  specific  characters  only  by  the  light  of 
analogy.  The  modifications  acquired  through  sexual  selection  are 
often  so  strongly  pronounced  that  the  two  sexes  have  frequently 
been  ranked  as  distinct  species,  or  even  as  distinct  genera.  Such 
strongly-marked  differences  must  be  in  some  manner  highly  im- 
portant; and  we  know  that  they  have  been  acquired  in  some  in- 
stances at  the  cost  not  only  of  inconvenience,  but  of  exposure 
to  actual  danger. 

The  belief  in  the  power  of  sexual  selection  rests  chiefly  on 
the  following  considerations.  Certain  characters  are  confined  to 
one  sex;  and  this  alone  renders  it  probable  that  in  most  cases 
they  are  connected  with  the  act  of  reproduction.  In  innumer- 
able instances  these  characters  are  fully  developed  only  at  ma- 
turity, and  often  during  only  a  part  of  the  year,  which  is  always 
the  breeding-season.  The  males  (passing  over  a  few  exceptional 
cases)  are  the  more  active  in  courtship;  they  are  the  better 
armed,  and  are  rendered  the  more  attractive  in  various  ways. 
It  is  to  be  especially  observed  that  the  males  display  their  at- 
tractions with  elaborate  care  in  the  presence  of  the  females;  and 
40 


610  THE   DESCENT  OF  MAN. 

that  they  rarely  or  never  display  them  excepting  during  the  sea- 
son of  love.  It  is  incredible  that  all  this  should  be  purposeless. 
Lastly,  we  have  distinct  evidence  with  some  quadrupeds  and 
birds,  that  the  individuals  of  one  sex  are  capable  of  feeling  a 
strong  antipathy  or  preference  for  certain  individuals  of  the  other 
sex. 

Bearing  in  mind  these  facts,  and  the  marked  results  of  man's 
unconscious  selection,  when  applied  to  domesticated  animals  and 
cultivated  plants,  it  seems  to  me  almost  certain  that  if  the  in- 
dividuals of  one  sex  were  during  a  long  series  of  generations  to 
prefer  pairing  with  certain  individuals  of  the  other  sex,  charac- 
terized in  some  peculiar  manner,  the  offspring  would  slowly  but 
surely  become  modified  in  this  same  manner.  I  have  not  at- 
tempted to  conceal  that,  excepting  when  the  males  are  more 
numerous  than  the  females,  or  when  polygamy  prevails,  it  is 
doubtful  how  the  more  attractive  males  succeed  in  leaving  a 
larger  number  of  offspring  to  inherit  their  superiority  in  orna- 
ments or  other  charms  than  the  less  attractive  males;  but  I 
have  shown  that  this  would  probably  follow  from  the  females, — 
especially  the  more  vigorous  ones,  which  would  be  the  first  to 
breed, — preferring  not  only  the  more  attractive  but  at  the  same 
time  the  more  vigorous  and  victorious  males. 

Although  we  have  some  positive  evidence  that  birds  appre- 
ciate bright  and  beautiful  objects,  as  with  the  bower-birds  of 
Australia,  and  although  they  certainly  appreciate  the  power  of 
song,  yet  I  fully  admit  that  it  is  astonishing  that  the  females 
of  many  birds  and  some  mammals  should  be  endowed  with  suf- 
ficient taste  to  appreciate  ornaments,  which  we  have  reason  to 
attribute  to  sexual  selection;  and  this  is  even  more  astonishing 
in  the  case  of  reptiles,  fish,  and  insects.  But  we  really  know 
little  about  the  minds  of  the  lower  animals.  It  cannot  be  sup- 
posed, for  instance,  that  male  birds  of  paradise  or  peacocks  should 
take  such  pains  in  erecting,  spreading,  and  vibrating  their  beauti- 
ful plumes  before  the  females  for  no  purpose.  We  should  re- 
member the  fact  given  on  excellent  authority  in  a  former  chapter, 
that  several  peahens,  when  debarred  from  an  admired  male,  re- 
mained widows  during  a  whole  season  rather  than  pair  with 
another  bird. 

Nevertheless  I  know  of  no  fact  in  natural  history  more  wonder- 
ful than  that  the  female  Argus  pheasant  should  appreciate  the 
exquisite  shading  of  the  ball-and-socket  ornaments  and  the  ele- 
gant patterns  on  the  wing-feathers  of  the  male.  He  who  thinks 
that  the  male  was  created  as  he  now  exists  must  admit  that 
the  great  plumes,  which  prevent  the  wings  from  being  used  for 
flight,  and  which  are  displayed  during  courtship  and  at  no  other 
time  in  a  manner  quite  peculiar  to  this  one  species,  were  given 
to  him  as  an  ornament.    If  so,  he  must  likewise  admit  that  the 


GENERAL  SUMMARY.  611 

female  was  created  and  endowed  with  the  capacity  of  appreciating 
8uch  ornaments.  I  differ  only  in  the  conviction  that  the  male 
Argus  pheasant  acquired  his  beauty  gradually,  through  the  pref- 
erence of  the  females  during  many  generations  for  the  more 
highly  ornamented  males;  the  aesthetic  capacity  of  the  females 
having  been  advanced  through  exercise  or  habit,  just  as  our  own 
taste  is  gradually  improved.  In  the  male  through  the  fortunate 
chance  of  a  few  feathers  being  left  unchanged,  we  can  distinctly 
trace  how  simple  spots  with  a  little  fulvous  shading  on  one  side 
may  have  been  developed  by  small  steps  into  the  wonderful  ball- 
and-socket  ornaments;  and  it  is  probable  that  they  were  actually 
thus  developed. 

Everyone  who  admits  the  principle  of  evolution,  and  yet  feels 
great  difficulty  in  admitting  that  female  mammals,  birds,  reptiles, 
and  fish,  could  have  acquired  the  high  taste  implied  by  the  beautj'' 
of  the  males,  and  which  generallj'-  coincides  with  our  own  stand- 
ard, should  reflect  that  the  nerve-cells  of  the  brain  in  the  highest 
as  well  as  in  the  lov/est  members  of  the  Vertebrate  series,  are 
derived  from  those  of  the  common  progenitor  of  this  great  King- 
dom. For  we  can  thus  see  how  it  has  come  to  pass  that  certain 
mental  faculties,  in  various  and  widely  distinct  groups  of  animals, 
have  been  developed  in  nearly  the  same  manner  and  to  nearly 
the  same  degree. 

The  reader  who  has  taken  the  trouble  to  go  through  the  several 
chapters  devoted  to  sexual  selection,  vv^ill  be  able  to  judge  how 
far  the  conclusions  at  which  I  have  arrived  are  supported  by 
sufficient  evidence.  If  he  accepts  these  conclusions  he  may,  I 
think,  safely  extend  them  to  mankind;  but  it  v/ould  be  super- 
fluous here  to  repeat  what  I  have  so  lately  said  on  the  manner  in 
which  sexual  selection  apparently  has  acted  on  man,  both  on  the 
male  and  female  side,  causing  the  two  sexes  to  differ  in  body  and 
mind,  and  the  several  races  to  differ  from  each  other  in  various 
characters,  as  v/ell  as  from  their  ancient  and  lowly-organized 
progenitors. 

He  who  admits  the  principle  of  sexual  selection  will  be  led 
to  the  remarkable  conclusion  that  the  nervous  system  not  only 
regulates  most  of  the  existing  functions  of  the  body,  but  has 
indirectly  influenced  the  progressive  development  of  various  bod- 
ily structures  and  of  certain  mental  qualities.  Courage,  pug- 
nacity, perseverance,  strength  and  size  of  body,  weapons  of  all 
kinds,  musical  organs,  both  vocal  and  instrumental,  bright  colors 
and  ornamental  appendages,  have  all  been  indirectly  gained  by 
the  one  sex  or  the  other,  through  the  exertion  of  choice,  the  in- 
fluence of  love  and  jealousy,  and  the  appreciation  of  the  beautiful 
in  sound,  color  or  form;  and  these  powers  of  the  mind  manifestly 
depend  on  the  development  of  the  brain. 


612  THE   DESCENT  OF  MAN. 

Man  scans  with  scrupulous  care  tlie  character  and  pedigree 
of  his  horses,  cattle  and  dogs  before  he  matches  them;  but 
when  he  comes  to  his  own  marriage  he  rarely,  or  never,  takes 
any  such  care.  He  is  impelled  by  nearly  the  same  motives  as 
the  lower  animals,  when  they  are  left  to  their  own  free  choice, 
though  he  is  in  so  far  superior  to  them  that  he  highly  values 
mental  charms  and  virtues.  On  the  other  hand  he  is  strongly 
attracted  by  mere  wealth  or  rank.  Yet  he  might  by  selection  do 
something  not  only  for  the  bodily  constitution  and  frame  of  his 
offspring,  but  for  their  intellectual  and  moral  qualities.  Both 
sexes  ought  to  refrain  from  marriage  if  they  are  in  any  marked 
degree  inferior  in  body  or  mind;  but  such  hopes  are  Utopian  and 
will  never  be  even  partially  realized  until  the  laws  of  inheritance 
are  thoroughly  known.  Everyone  does  good  service,  who  aids 
towards  this  end.  When  the  principles  of  breeding  and  inheri- 
tance are  better  understood,  we  shall  not  hear  ignorant  members 
of  our  legislature  rejecting  with  scorn  a  plan  for  ascertaining 
whether  or  not  consanguineous  marriages  are  injurious  to  man. 

The  advancement  of  the  welfare  of  mankind  is  a  most  intricate 
problem:  all  ought  to  refrain  from  marriage  who  cannot  avoid 
abject  poverty  for  their  children;  for  poverty  is  not  only  a  great 
evil,  but  tends  to  its  own  increase  by  leading  to  recklessness  in 
marriage.  On  the  other  hand,  as  Mr.  Galton  has  remarked,  if 
the  prudent  avoid  marriage,  whilst  the  reckless  marry,  the  in- 
ferior members  tend  to  supplant  the  better  members  of  society. 
Man,  like  every  other  animal,  has  no  doubt  advanced  to  his  pres- 
ent high  condition  through  a  struggle  for  existence  consequent 
on  his  rapid  multiplication;  and  if  he  is  to  advance  still  higher, 
it  is  to  be  feared  that  he  must  remain  subject  to  a  severe  struggle. 
Otherwise  he  would  sink  into  indolence,  and  the  more  gifted 
men  would  not  be  more  successful  in  the  battle  of  life  than  the 
less  gifted.  Hence  our  natural  rate  of  increase,  though  leading 
to  many  and  obvious  evils,  must  not  be  greatly  diminished  by 
any  means.  There  should  be  open  competition  for  all  men;  and 
the  most  able  should  not  be  prevented  by  laws  or  customs  from 
succeeding  best  and  rearing  the  largest  number  of  offspring.  Im- 
portant as  the  struggle  for  existence  has  been  and  even  still  is, 
yet  as  far  as  the  highest  part  of  man's  nature  is  concerned  there 
are  other  agencies  more  important.  For  the  moral  qualities  are 
advanced,  either  directly  or  indirectly,  much  more  through  the 
effects  of  habit,  the  reasoning  powers,  instruction,  religion,  &c., 
than  through  natural  selection;  though  to  this  latter  agency  may 
be  safely  attributed  the  social  instincts,  which  afforded  the  basis 
for  the  development  of  the  moral  sense. 

The   main  conclusion  arrived   at   in  this   work,   namely  that 
%ian  is  descended  from  some  lowly  organized  form,  will,  I  regret 


GENERAL  SUMMARY.  613 

to  think,  be  highly  distasteful  to  many.  But  there  can  hardly 
be  a  doubt  that  we  are  descended  from  barbarians.  The  aston- 
ishment which  I  felt  on  first  seeing  a  party  of  Fuegians  on  a 
wild  and  broken  shore  will  never  be  forgotten  by  me,  for  the 
reflection  at  once  rushed  into  my  mind — such  were  our  ancestors. 
These  men  were  absolutely  naked  and  bedaubed  with  paint,  their 
long  hair  was  tangled,  their  mouths  frothed  with  excitement, 
and  their  expression  was  wild,  startled,  and  distrustful.  They  pos- 
sessed hardly  any  arts,  and  like  wild  animals  lived  on  what  they 
could  catch;  they  had  no  government,  and  were  merciless  to 
every  one  not  of  their  own  small  tribe.  He  who  has  seen  a  sav- 
age in  his  native  land  will  not  feel  much  shame,  if  forced  to  ac- 
knowledge that  the  blood  of  some  more  humble  creature  flows 
in  his  veins.  For  my  own  part  I  would  as  soon  be  descended  from 
that  heroic  little  monkey,  who  braved  his  dreaded  enemy  in  order 
to  save  the  life  of  his  keeper,  or  from  that  old  baboon,  who  de- 
scending from  the  mountains,  carried  away  in  triumph  his  young 
comrade  from  a  crowd  of  astonished  dogs — as  from  a  savage  who 
delights  to  torture  his  enemies,  offers  up  bloody  sacrifices,  prac- 
tices infanticide  without  remorse,  treats  his  wives  like  slaves, 
knows  no  decency,  and  is  haunted  by  the  grossest  superstitions. 

Man  may  be  excused  for  feeling  some  pride  at  having  risen, 
though  not  through  his  own  exertions,  to  the  very  summit  of  the 
organic  scale;  and  the  fact  of  his  having  thus  risen,  instead  of 
having  been  aboriginally  placed  there,  may  give  him  hope  for  a 
still  higher  destiny  in  the  distant  future.  But  we  are  not  here 
concerned  with  hopes  or  fears,  only  with  the  truth  as  far  as  our 
reason  permits  us  to  discover  it;  and  I  have  given  the  evidence 
to  the  best  of  my  ability.  We  must,  however,  acknowledge,  as  it 
seems  to  me,  that  man  with  all  his  noble  qualities,  with  sympathy 
which  feels  for  the  most  debased,  with  benevolence  which  extends 
not  only  to  other  men  but  to  the  humblest  living  creature,  with 
his  god-like  intellect  v/hich  has  penetrated  into  the  movements 
and  constitution  of  the  solar  system — with  all  these  exalted  pow- 
ers— Man  still  bears  in  his  bodily  frame  the  indelible  stamp  of  his 
lowly   origin. 


THE  END. 


kO 


INDEX. 


Abbot,  C,  on  the  battles  of  seals, 
496. 

Abductor  of  the  fifth  metatarsal, 
presence  of,  in  man,  40. 

Abercrombie,  Dr.,  on  disease  of 
the  brain  affecting  speech,  86. 

Abipones,  marriage  customs  of 
the,  592. 

Abou-Simbel,    caves  of,   164. 

Abortion,  prevalence  of  the  prac- 
tice of,  45. 

Abstraction,  power  of,  in  ani- 
mals, 80. 

Acalles,  stridulation  of,  303. 

Acanthodactylus  capensis,  sex- 
ual  differences    of   color  in,    353. 

Accentor  modularis,  467. 

Acclimatization,  difference  of,  in 
different  races   of  men,  163. 

Achetidae,  stridulation  of  the, 
280,  283;  rudimentary  stridulat- 
ing  organs   in   female,  285. 

Acilius  sulcatus,  elytra  of  the 
female,  273. 

Acomus,  development  of  spurs  in 
the  female  of,  444. 

Acridiidae,  stridulation  of  the, 
280,  283;  rudimentary  stridulat- 
ing  organs  in  female,   285. 

Acromio-basilar  m.uscle,  and 
quadrupedal   gait,    40. 

Acting,    174. 

Actiniae,  bright  colors  of,  257. 

Adams,  Mr.,  migration  of  birds, 
105;  intelligence  of  nut-hatch, 
405;  on  the  Bombycilla  carolin- 
Giisis    455 

Admiral  butterfly,  309. 

Adoption  of  the  young  of  other 
animals  by  female  monkeys,  68. 

Advancement  in  the  organic 
scale,  Von  Baer's  definition  of, 
159. 

JEby,  on  the  difference  between 
the  skulls  of  man  and  the 
quadrumana,   145. 

^slhetic  faculty,  not  highly  de- 
■V  eloped  in  savages,  90. 

Affection,  maternal,  68;  manifes- 
tation of,  by  animals,  68;  par- 
ental and  filial,  partly  the  re- 
sult of  natural  selection,  102; 
mutual,  of  birds,  405;    shown  by 


615 


birds  in  confinement,  for  cer- 
tain persons,  406. 
Africa,  probably  the  birthplace 
of  man,  151;  South  crossed 
population  of,  170;  South,  re- 
tention of  color  by  the  Dutch 
in,  188;  South,  proportion  of  the 
sexes  in  the  butterflies  of,  246; 
tattooing  practiced  in,  569; 
Northern,  coiffure  of  natives 
of,  569. 

Agassiz,  L..,  on  conscience  in 
dogs,  100;  on  the  coincidence  of 
the  races  of  man  with  zoologi- 
cal provinces,  165;  on  the  num- 
ber of  species  of  man,  170;  on 
the  courtship  of  the  land- 
snails,  259;  on  the  brightness  of 
the  colors  of  male  fishes  during 
the  breeding  season,  336:  on 
the  frontal  protuberance  of  the 
males  of  Geophagus  and  Cich- 
la,  336,  341;  male  fishes  hatching 
ova  in  their  mouths,  342:  sexual 
differences  in  color  of  chro- 
mids,  342;  on  the  slight  sexual 
differences  of  the  South  Ameri- 
cans, 556;  on  the  tattooing  of 
the  Amazonian  Indians,  571. 

Age,  in  relation  to  the  transmis- 
sion of  characters  in  birds,  458; 
variation  in  accordance  with, 
in  birds,  478. 

Agelaeus  phoeniceus,  220,  410. 

Ageronia  feronia,  noise  produced 
by,  304. 

Agrion,   dimorphism  in,   288. 

Ramburii,   sexes  of,  288. 

Agrionidse,  difference  in  the  sex- 
es of,  288. 

Agrotis  exclamationis,  313. 

Ague,  tertian,  dog,  suffering 
from,   8. 

Aithurus  polytmus,  young  of, 
483. 

Ainos,  hairiness  of  the,  555. 

Albino  birds,  413. 

Alca  torda,  young  of,  481. 

A-lces  palmata,  511. 

Alder  and  Hancock,  MM.,  on  the 
nudi-branch  mollusca,  261. 

Allen,  J.  A.,  vigor  of  birds  ear- 
liest hatched,  208,  209;  effect  of 
difference  of  temperature, 
light,   &c.,  on  birds,  220;    colors 


616 


INDEX. 


of  birds,  416;  on  the  relative 
size  of  the  sexes  of  Callorhinus 
ursinus,  511;  on  the  mane  of 
Otaria  jubata,  516;  on  the  pair- 
ing- of  seals,  518;  on  sexual  dif- 
ferences in  the  color  of  bats, 
529. 

Allen,  S.,  on  the  habits  of  Hoplo- 
pterus,  362;  on  the  plumes  of 
herons,  386;  on  the  vernal  moult 
of  Herodias  bubulcus,  387. 

Alligator,  courtship  of  the  male, 
216,  347;  roaring  of  the  male, 
561. 

Amadavat,     pugnacity     of     male, 

362.  ,.     , 

Amadina  Lathami,  display  of 
plumage  by  the  male,   396. 

castanotis,  display  of  plum- 
age by  the  male,  396. 

Amazons,  butterflies  of  the,  246; 
fishes  of  the,  339. 

America,  variation  in  the  skulls 
of  aborigines  of,  25;  wide  range 
of  aborigines  of,  165;  lice  of  the 
natives  of,  166;  general  beard- 
lessness  of  the  natives  of,  555. 

,  North,     butterflies    of,     246; 

Indians  of,  women  a  cause  of 
strife  among  the,  556;  Indians 
of,  their  notions  of  female 
beauty,  571,  572. 

•,  South,  character  of  the  na- 


tives of,  164;  population  of 
parts  of,  169;  piles  of  stone  in, 
175;  extinction  of  the  fossil 
horse  of,  186;  desert-birds  of, 
485;  slight  sexual  difference  of 
the  aborigines  of,  556;  preva- 
lence of  infanticide  in,  586. 

American  languages,  often  high- 
ly artificial,  88. 

Americans,  wide  geographical 
range  of,  28;  native,  variability 
of  170;  and  negroes,  difCerence 
of,  192;  aversion  of,  to  hair  on 
the    face,    574. 

Ammophila,  on  the  jaws  of,  273. 

Ammotragus  tragelaphus,  hairy 
forelegs  of,  526,  528. 

Amphibia,  affinity  of,  to  the  gan- 
oid fishes,  154;  vocal  organs  of 
the,  561. 

Amphibians,  160,  344;  breeding 
whilst  Immature,   480. 

Amphioxus,  155. 

Amphipoda,  males  sexually  ma- 
ture while  young,  480. 

Amunoph  III.,  negro  character 
of  features  of,  164. 

Anal  appendages  of  insects,  272-3. 

Analogous  variation  in  the  plum- 
age of  birds,   880. 

A  Y^QC!      456 

'acuta,    male     plumage      of, 

387. 
. boschas,   male  plumage    of, 

387. 

histrionica,  479. 

'■ punctata,    369. 


Anastomus  oscitans,  sexes  and 
young  of,  481;  white  nuptial 
plumage  of,   48S. 

Anatidae,  voices  of,  370. 

Anax  Junius,  differences  in  the 
sexes  of,   287. 

Andaman  islanders,  susceptible 
to  change  of  climate,  183. 

Anderson,  Dr.,  on  the  tail  of 
Macacus  brunneus,  57;  the  Bu- 
fo  sikimmensis,  345;  sounds  of 
Echis  carinata,   349. 

Andrsena  fulva,  290. 

Anglo-Saxons,  estimation  of  the 
beard  among   the,   575. 

Animals,  domesticated,  more  fer- 
tile than  wild,  43;  cruelty  of 
savages  to,  114;  characters  com- 
mon to  man  and,  142;  domestic, 
change  of  breeds  of.  590. 

Annelida,  261;    colors  of,  261. 

Anobium  tessellatum,  sounds 
produced    by,    302. 

Anolis  cristatellus,  male,  crest 
of,  350;  pugnacity  of  the  male, 
349;    throat-pouch  of,  350. 

Anser  canadensis,   411. 

cygnoides,  409;    knob  at  the 

base  of  the  beak  of,  420. 

hyperboreus,    whiteness   of. 


Antelope,  prong-horned,  horns 
of,  229. 

Antelopes,  generally  polygamous, 
213;  horns  of,  229,  500;  canine 
teeth  of  some  male,  498;  use  of 
horns  of,  504;  dorsal  crests  in, 
526;  dewlaps  of,  527;  winter 
change  of  two  species  of,  538; 
peculiar  markings  of,   538. 

Antennae,  furnished  with  cush- 
ions in  the  male  of  Pentha,  273. 

Anthidium  manicatum,  large 
male  of,    277. 

Anthocharis  cardamines,  306,  30&; 
sexual  difference  of  color  in, 
319. 

genutia,    309. 

sara,   309. 

Anthophora  acervorum,  large 
male   of,   277. 

retusa,      difference    of     the 

sexes  in,  290. 

Anthropidse,  148. 

Anthus,  moulting  of,  386. 

Antics  of  birds,   375. 

Antigua,  Dr.  Nicholson's  obser- 
vations on  yellow  fever  in,   190. 

Antilocapra  americana,  horns  of, 
229,  500,  503. 

Antilope  bezoartica,  horned  fe- 
males of,  500,  502,  503;  Sexual 
difference  in  the  color  of,  531. 

Dorcas  and  euchore,  500. 

euchore,  horns  of,   504. 

montana,  rudimentary  ca- 
nines in  the  young  male  of,  510. 

niger,     sing     sing,       caama, 

and  gorgon,  sexual  difference? 
in  the  colors  of,  531. 


INDEX. 


617 


Antilope  oreas,  horns  of,  229. 

saiga,      polygamous     habits 

of,  213. 

strepsiceros,    horns    of,    229. 

subgutturosa,      absence      of 

suborbital  pits  in,  525. 

Antipathy,  shown  by  birds  in 
confinement,  to  certain  per- 
sons, 406. 

Ants,  143;  large  size  of  the  cere- 
bral ganglia  in,  52;  soldier, 
large  jaws  of,  60;  playing  to- 
gether, 67;  memory  in,  72;  inter- 
communication of,  by  means  of 
the  antennae,  87;  habits  of, 
143;  difference  of  the  sexes  in, 
289;  recognition  of  each  other 
by,  after  separation,  289. 

■ White,  habits  of,  288. 

Anura,  345. 

Apatania  muliebris,  male  un- 
known, 251. 

Apathus,  difference  of  the  sexes 
in,  290. 

Apatura  Iris,  304,  305, 

Apes,  difference  of  the  young, 
from  the  adult,  8;  semi-erect 
attitude  of  some,  50;  mastoid 
processes  of,  51;  influences  of 
the  jaw-muscles  on  the  physi- 
ognomy of,  52;  female  destitute 
of  large  canines,  60;  building 
platforms,  80;  imitative  facul- 
ties of,  126;  anthropomorphous, 
149;  probable  speedy  extermin- 
ation of  the,  152;  Gratiolet  on 
the  evolution  of,  173;  canine 
teeth  of  male,  498;  females  of 
some,  less  hairy  beneath  than 
the  males,  595. 

long-armed,    their    mode    of 

progression,  50. 

Aphasia,  Dr.  Bateman  on,  86. 

Apis  melliflca,  large  male  of,  277. 

Apollo,   Greek  statues  of,  576. 

Apoplexy  in  Cebus  Azarse,  7. 

Appendages,  anal,  of  insects, 
274,  275. 

Approbation,  influence  of  the 
love  of,  106,  112,  128. 

Aprosmictus  scapulatus,  451. 

Apus,  proportion  of  sexes,  251. 

Aquatic  birds,  frequency  of 
white   plumage   in,   490. 

Aqulla  chryssetos,  403. 

Arab  women,  elaborate  and  pe- 
culiar coiffure  of,  578, 

Arabs,  fertility  of  crosses  with 
other  races,  167;  gashing  of 
cheeks  and  temples  among  the, 
569. 

Arachnida,    268. 

Arakhan,  artificial  widening  of 
the  forehead  by  the  natives  of, 
577. 

Arboricola,  young  of,  463. 

Archeopteryx,    154. 

Arctiidse,     coloration    of    the,  311. 

Ardea  asha,  rufescens,  and  c£er- 
ulea,  ehange  of  color  in,  490,  491. 


Ardea    cserulea,    breeding    in    im- 
mature plumage,  479. 

gularis,   change   of   plumage 

in,  490. 

herodias,       love-gestures    of 

the  male,  375. 

ludoviciana,   age  of  mature 

plumage  in,  478;  continued 
growth  of  crest  and  plumes  in 
the  male  of,  480, 

nycticorax,   cries  of,   363. 


Ardeola,  young  of,   463. 

Ardetta,  changes  of  plumage  in, 
454. 

Argenteuil,  21,  22. 

Argus  pheasant,  379,  397,  456;  dis- 
play of  plumage  by  the  male, 
394;  ocellated  spots  of  the,  422; 
gradation  of  characters  in  the, 
428. 

Argyll,  Duke  of,  on  the  physi- 
cal weakness  of  man,  60;  the 
fashioning  of  implements  pe- 
culiar to  man,  80;  on  the  con- 
test in  man  between  right  and 
wrong,  121;  on  the  primitive 
civilization  of  man,  140;  on  the 
plum.age  of  the  male  Argus 
pheasant,  392;  on  Urosticte 
Benjamini,  436,  437;  on  the  nests 
of  birds,   447. 

Argynnis,  coloring  of  the  lower 
surface   of,   311. 

Aricoris  epitus,  sexual  differ- 
ences in  the  wings  of,  274. 

Aristocracy,  increased  beauty  of 
the,  581. 

Arms,  proportions  of,  in  soldiers 
and  sailors,  31;  direction  of  the 
hair  on  the,  147. 

and  hands,  free  use  of,  in- 
directly correlated  with  dimin- 
ution of  canines,  51. 

Arrest  of  development,  34,  35. 

Arrow-heads,  stone,  general  re- 
semblance of,  174. 

Arrows,  use  of,  174. 

Arteries,  variations  in  the  course 
of  the,  25, 

Artery,  effect  of  tying,  upon  the 
lateral  channels,  31. 

Arthropoda,    262. 

Arts  practiced  by  savages,  174. 

Ascension,  colored  incrustation 
on  the  rocks  of,  260. 

Ascidia,  affinity  of  the  lancelet 
to,  155;  tadpole-like  larvae  of, 
155. 

Ascidians,  260;  bright  colors  of 
some,  257. 

Asinus,  Asiatic  and  African  spe- 
cies of,  543. 

tsaniopus,  543, 

Ass,    color-variations    of    the,    542, 
Ateles,    effects   of   brandy   on    an, 
7;    absence   of   the  thumb  in,  49. 
beelzebuth,  ears  cf,  15. 

marginatus,     color     of     the 

ruff   of,    532;     hair  on   the    head 
of.  544. 


618 


INDEX. 


Ateuchus,  stridulation  of,  303. 

,  cicatricosus,  habits  of,  297. 

Athalia,  proportions  of  the  sexes 
in,   250. 

Atropus  pulsatorius,  288, 

Attention,  manifestations  of,  in 
animals,  71. 

Audouin,  V.,  on  a  hymenopterous 
parasite  with  a  sedentary  male, 
217. 

Auduhon,  J.  J.,  on  the  pinioned 
goose,  101;  on  the  speculum  of 
Mergus  cucullatus,  231;  on  the 
pugnacity  of  male  birds,  358, 
362;  on  Tetrao  cupido,  363;  on 
Ardea  nycticorax,  363;  on  Stur- 
nella  ludoviciana,  363;  on  the 
vocal  organs  of  Tetrao  cupido, 
367;  on  the  drumming  of  the 
male  Tetrao  umbellus,  371;  on 
sounds  produced  by  the  night- 
jar 371;  on  Ardea  herodias  and 
Cathartes  jota,  375;  on  Mimus 
polyglottus,  406;  on  display  m 
male  birds,  389;  on  the  spring 
change  of  color  in  some  finches, 
388;  recognition  of  a  dog  by  a 
turkey,  407;  selection  of  mate 
by  female  birds,  411;  on  the  tur- 
key 406,  414;  on  variation 
in  'the  male  scarlet  tanager, 
418-  on  the  musk-rat,  537;  on  the 
habits  of  Pyranga  estiva,  447; 
on  local  differences  in  the  nests 
of  the  same  species  of  birds, 
450-  on  the  habits  of  wood- 
peckers, 452;  on  Bombycilla 
carolinensis,  455;  on  young  fe- 
males of  Tyranga  sestiva  ac- 
quiring male  characters,  455; 
on  the  immature  plum.age  of 
thrushes,  459;  on  the  immature 
plumage  of  birds,  459,  et  seq.; 
on  birds  breeding  in  immature 
plumage,  479;  on  the  growth  of 
the  crest  and  plumes  m  the 
male  Ardea  ludoviciana,  480;  on 
the  change  of  color  in  some 
species  of  Ardea,  490. 
Audubon  and  Bachman,  M.M.,  on 
squirrels  fighting,  496;  on  the 
Canadian  lynx,  516. 
Aughey,    Prof.,    on     rattlesnaxiies, 

349. 
Austen,  N.  I^.,  on  Anolis  cristat- 
ellus,  350.  ,  .  .^   ,  f 

Australia,  not  the  birthplace  of 
man  151;  half-castes  killed  by 
the  natives  of,  166;  lice  of  the 
natives  of,  166;  prevalence  of  fe- 
male infanticide  in,  586. 

,     South,      variation     in    the 

skulls  of  aborigines  of,  25. 
Australians,  cclcr  of  newborn 
children  of,  552;  relative  height 
of  the  sexes  of,  554;  women  a 
cause  of  war  among  the,  556. 
Axis  deer,  sexual  difference  in 
the  color  of  the,  532. 


Ayraaras,  measurements  of  the, 
33;  no  gray  hair  among  the, 
554;  hairlessness  of  the  face  in 
the,  555;    long  hair  of  the,  574. 

Azara,  on  the  proportion  of  men 
and  women  among  the  Guar- 
anys,  238;  on  Palamedea  cor- 
nuta,  361;  on  the  beards  of  the 
Guaranys,  556;  on  strife  for 
women  among  the  Guanas,  556; 
on  infanticide,  572,  587;  on  the 
eradication  of  the  eyebrows 
and  eyelashes  by  the  Indians 
of  Paraguay,  575;  on  polyandry 
among  the  Guanas,  588;  celi- 
bacy unknown  among  the  sav- 
ages of  South  America,  588;  on 
the  freedom  of  divorce  among 
the  Charruas,   592. 


B 

Babbage,  C,  on  the  greater  pro- 
portion of  illegitimate  female 
births,  239. 

Babirusa,  tusks  of  the,  515. 

Baboon,  revenge  in  a,  67;  rage 
excited  in,  by  reading,  69;  man- 
ifestation of  memory  by  a,  72; 
employing  a  mat  for  shelter 
against  the  sun,  80;  protected 
from  punishment  by  its  com- 
panions,  100. 

,     Cape,    mane   of   the  male, 

516;     Hamadryas,    mane    of    the 
male,  517. 

Baboons,    effects    of    intoxicating 
liquors  on,  7;    ears  of,  15;    diver- 
sity of  the  mental  faculties  in 
26;    hands  of,    48;     habits  of,   49 
variability   of     the     tail     in,    56 
manifestation    of    m.aternal    af- 
fection by,  68;    using  stones  and 
sticks  as  weapons,  79;    co-opera- 
tion of,  97;    silence  of,  on  plun- 
dering expenditions,  101;    appar- 
ent polygamy    of,     212;     polyga- 
mous  and  social  habits    of.    585. 

Bachman,  Dr.,  on  the  fertility  of 
mulattoes,  167. 

Baer,  K.  E.,  von,  on  embrj'-onic 
development,  9. 

Bagehot,    W.,    on   the    social   vir- 
tues  among  primitive  men,   113 
slavery  formerly  beneficial,   114 
on   the   value   of  obedience,  127 
on  human  progress,  129;    on  the 
persistence   of   savage   tribes    in 
classical  times,   179. 

Bailly,  E.  M.,  on  the  mode  of 
fighting  of  the  Italian  buffalo, 
504;     on    the    fighting    of    stags, 

Bain,  A.,  on  the  sense  of  duty, 
95;  aid  springing  from  sympa- 
thy. 100;  on  the  basis  of  sym- 
pathy, 103;  on  love  of  approba- 
tion, &c.,  106;  on  the  idea  Ox 
beauty,  579. 


INDEX. 


619 


feaird,  W.,  on  a  difference  in 
color  between  the  males  and  fe- 
males of  some  Entozoa,  257. 

Baker,  Mr.,  observation  on  the 
proportion  of  the  sexes  in 
pheasant-chicks,   242. 

,    Sir  S.,    on  the   fondness  of 

the  Arabs  for  discordant  music, 
375;  on  sexual  difference  in  the 
colors  of  an  antelope,  531;  on 
the  elephant  and  rhinoceros  at- 
tacking white  or  gray  horses, 
535;  on  the  disfigurements  prac- 
ticed by  the  negroes,  536;  on 
the  gashing  of  the  cheeks  and 
temples  practiced  in  Arab  coun- 
tries, 569;  on  the  coiffure  of  the 
North  Africans,  569;  on  the  per- 
foration of  the  lower  lip  by 
the  women  of  Latooka,  570;  on 
the  distinctive  characters  of 
the  coiffure  of  central  African 
tribes,  571;  on  the  coiffure  of 
Arab  women,   578. 

"Balz"  of  the  Black-cock,  359,  400. 

Bantam,  Sebright,  207,  233. 

Banteng,  horns  of,  501;  sexual 
differences  in  the  colors  of  the, 
531. 

Banyai,  color  of  the,  573. 

Barbarism,  primitive,  of  civilized 
nations,  140. 

Barbs,  filamentous,  of  the  feath- 
ers, in  certain  birds,  380,  424. 

Barrago,  F.,  on  the  Simian  re- 
semblances of  man,  2. 

Barr,  Mr.,  on  sexual  preference 
in  dogs,  520. 

Barrington,  Daines,  on  the  lan- 
guage of  birds,  84;  on  the  cluck- 
ing of  the  hen,  363;  on  the  ob- 
ject of  the  song  of  birds,  364; 
on  the  singing  of  female  birds, 
S65;  on  birds  acquiring  the 
songs  of  other  birds,  365;  on  the 
muscles  of  the  larynx  in  song 
birds,  366;  on  the  want  of  the 
power  of  song  by  female  birds, 
444. 

Barrow,  on  the  widow-bird,  387. 

Bartels,  Dr.,  supernumerary 
mammse  in  men,   36. 

Bartlett,  A.  D.,  period  of  hatch- 
ing of  birds'  eggs,  161;  on  the 
tragopan,  215;  on  the  develop- 
ment of  the  spurs  in  Crossop- 
tilon  auritum.,  230;  on  the  fight- 
ing of  the  males  of  Plectrcpter- 
us  gambensis,  360;  on  the  knot, 
386;  on  display  in  male  birds, 
389;  on  the  displaj'-  of  phimage 
by  the  male  Polyplectron,  391; 
on  Crossoptilon  auritum  and 
Phasianus  Wallichii,  395;  on  the 
habits  of  Dophophorus,  414;  on 
the  color  of  the  mouth  in  Bucer- 
cs  bicornis,  420;  on  the  incuba- 
tion of  the  cassowary,  472;  on 
the  Cape  buffalo,  504;  on  the 
use   of  the   horns   of   antelopes, 


505;  on  the  fighting  of  male 
wart-hogs,  516;  on  Ammotragus 
tragelaphus,  526;  on  the  colors  of 
Cercopithecus  cephus,  532;  on 
the  colors  of  the  faces  of 
monkeys,  546;  on  the  naked  sur- 
faces of  monkeys,  595. 

Bartram,  on  the  courtship  of  the 
male    alligator,    347. 

Basque  language,  highly  artifi- 
cial, 88. 

Bate,  C.  S.,  on  the  superior  activ- 
ity of  male  Crustacea,  217;  on  the 
proportions  of  the  sexes  in 
crabs,  252;  on  the  chelae  of 
Crustacea,  263;  on  the  relative 
size  of  the  sexes  in  Crustacea, 
265;  on  the  colors  of  Crustacea, 
267. 

Eatem.an,  Dr.,  tendency  to  imita- 
tion in  certain  diseased  states, 
71;    on  Aphasia,  86. 

Bates,  H.  W.,  on  variation  in  the 
form  of  the  head  of  Amazonian 
Indians,  27;  on  the  proportion  of 
the  sexes  among  Amazonian 
butterflies,  246;  on  sexual  differ- 
ences in  the  wings  of  butterflies, 
274;  on  the  fleld-cricket,  280;  on 
Pyrodes  pulcherrimus,  291;  on 
the  horns  of  Lamellicorn  bee- 
tles, 293,  294;  on  the  colors  of 
Epicaliee,  «6;c.,  306;  on  the  color- 
ation of  tropical  butterflies,  308; 
on  the  variability  of  Papilio  Se- 
sostris  and  Childrenee,  316;  on 
male  and  female  butterflies  in- 
habiting different  stations,  317; 
on  mimicry,  319;  on  the  cater- 
pillar of  a  Sphinx,  321;  on  the 
vocal  organs  of  the  umbrella- 
bird,  368;  on  the  toucans,  487;  on 
Brachyurus  calvus,  546. 

Batokas,  knocking  out  two  upper 
incisors,    569. 

Batrachia,  345;  eagerness  of  male, 
216. 

Bats,  scent-glands,  524;  sexual  dif- 
ferences in  the  color  of,  529;  fur 
of  male  frugivorous,  529. 

Battle,  law  of,  141;  among  bee- 
tles, 296;  among  birds,  356; 
among  mammals,  496  et  seq. ;  in 
man,  556. 

Beak,  sexual  difference  in  the 
forms  of  the,  355;  in  the  color  of 
the,   377. 

Beaks,  of  birds,  bright  colors  of, 

487. 

Beard,  development  of,  in  man, 
552;  analogy  of  the,  in  man  and 
the  quadrumana,  553;  variation 
of  the  development  of  the,  in 
different  races  of  men,  554;  es- 
timation of,  among  bearded  na- 
tions, 575;  probable  origin  of  the, 
597. 

,   in  monkeys,   146;    of  mam* 

mals,  526. 


620 


INDEX. 


Beautiful,  taste  for  the,  in  birds, 
405;    in  the  quadrumana,  535. 

Beauty,  sense  of,  in  animals,  89; 
appreciation  of,  by  birds,  408; 
influence  of,  568,  570;  variability 
of  the  standard  of,  591. 

Beavan,  Lieut.,  on  the  develop- 
ment of  the  horns  in  Cervus 
Eldi    229. 

Beaver,  instinct  and  intelligence 
of  the,  65,  66;  voice  of  the,  523; 
castoreum  of  the,  524. 

Beavers,  battles  of  male,  496. 

Bechstein,  on  female  birds  choos- 
ing the  best  singers  among  the 
males,  364;  on  rivalry  in  song- 
birds, 364;  on  the  singing  of  fe- 
male birds,  365;  on  birds  acquir- 
ing the  songs  of  other  birds,  366; 
on  pairing  the  canary  and  siskm, 
410;  on  a  subvariety  of  the  monk 
pigeon,  421;  on  spurred  hens,  443. 

Beddoe,  Dr.,  on  causes  of  differ- 
ence in  stature,  30. 

Bee-eater,    366. 

Bees,  96;  pollen-baskets  and  stings 
of,  60;  destruction  of  drones  and 
queens  by,  102;  female,  secondary 
sexual  characters  of,  204;  pro- 
portion of  sexes,  250;  difference 
of  the  sexes  in  color  and  sexual 
selection,  289. 

Beetle,  luminous  larva  of  a,  275. 

Beetles,  291;  size  of  the  cerebral 
ganglia  in,  52;  dilatation  of  the 
fore  tarsi  in  male,  273;  blind,  291; 
stridulation  of,   298. 

Belgium,  ancient  Inhabitants  of, 
177. 

Bell,  Sir  C,  on  emotional  muscles 
in  man,  3;  "snarling  muscles," 
39;    on  the  hand,  49. 

,  T.,  on  the  numerical  pro- 
portion of  the  sexes  in  moles, 
242;  on  the  newts,  344;  on  the 
croaking  of  the  frog,  346;  on  the 
difference  in  the  coloration  of 
the  sexes  in  Zootoca  vivipara, 
853;    on  moles  fighting,  496. 

Bell-bird,  sexual  difference  in  the 
color  of  the,   384. 

Bell-birds,   colors   of,  488. 

Belt,  Mr.,  on  the  nakedness  of 
tropical  mankind,  55;  on  a 
spider-monkey  and  eagle,  98; 
habits  of  ants,  143;  Lampyridse 
distasteful  to  mammals,  275; 
mimicry  of  Leptalides,  321;  col- 
ors of  Nicaraguan  frogs,  345; 
display  of  humming-birds,  437; 
on  the  toucans,  487;  protective 
coloring  of  skunk,   538. 

Benevolence,  manifested  by  birds, 
406. 

Bennett,  A.  W.,  attachment  of 
mated  birds,  405;  on  the  habits 
of  Dromceus  irroratus,  473. 

,    Dr.,    on  birds   of  paradise, 

390. 


Berbers,  fertility  of  crosses  with 
other  races.   167. 

Bernicla  antarctica,  colors  of,  488. 

Beinicle  gander  pairing  with  a 
Canada  goose,  409. 

Bert,  M.,  crustaceans  distinguish 
colors,   267. 

Bertillon,  M.,  arrested  develop- 
ment and  polydactylism,  35. 

Bettoni,  E.,  on  local  differences  in 
the  nests  of  Italian  birds,  450. 

Bevle,  M.,  see  Bombet. 

Bhoteas,  color  of  the  beard  in, 
553. 

Bhringa,  disciform  tail-feathers 
of,    387. 

Bianconi,  Prof.,  on  structures  as 
explained  through  mechanical 
principles,  23. 

Bibio,  sexual  differences  in  the 
genus,  277. 

Bichat,  on  beauty,  579. 

Bickes,  proportion  of  sexes  in 
man,    237. 

Bile,  colored,  in  many  animals, 
258. 

Bimana,  145. 

Birds,  imitations  of  the  songs  of 
other  birds  by,  71;  dreaming,  72; 
killed  by  telegraph  wires,  78; 
language  of,  83;  sense  of  beauty 
in,  89;  pleasure  of,  in  incubation, 
101;  male.  Incubation  by,  158;  and 
reptiles,  alliance  of,  160;  sexual 
differences  in  the  beak  of  some, 
204;  migratory,  arrival  of  the 
male  before  the  female,  208;  ap- 
parent relation  between  polyg- 
amy and  marked  sexual  differ- 
ences in,  215;  monogamous,  be- 
coming polygamous  under  do- 
mestication, 215;  eagerness  of 
male  in  pursuit  of  the  female, 
216;  wild,  numerical  proportion 
of  the  sexes  in,  242;  secondary 
sexual  characters  of,  355;  differ- 
ence of  size  in  the  sexes  of,  358; 
fights  of  male,  witnessed  by  fe- 
males, 362;  display  of  male,  to 
captivate  the  females,  363;  close 
attention  of,  to  the  songs  of  oth- 
ers, 364;  acquiring  the  song  of 
their  foster-parents,  366;  bril- 
liant, rarely  good  songsters,  366; 
love-antics  and  dances  of,  375; 
coloration  of,  380  et  seq.; 
moulting  of,  390  et  seq.;  un- 
paired, 401;  male,  singing  out  of 
season,  404;  mutual  affection  of, 
405;  in  confinement,  distinguish 
persons,  406;  hybrid,  production 
of,  408;  Albino,  413;  European, 
number  of  species  of,  416;  varia- 
bility of,  416;  geographical  dis- 
tribution of  coloring,  416;  grada- 
tion of  secondary  sexual  char- 
acters in,  424;  obscurely  colored, 
building  concealed  nests,  448; 
young  female  acquiring  male 
characters,  455;    breeding  in  im- 


INDEX. 


621 


mature  plumage,   479;    moulting 
of,    478;     aquatic    frequency    of 
white    plumage    in,     488;      vocal 
courtship  of,  562;    naked  skin  of 
the  head  and  neck  in,  595. 
Birgus  latro,  habits  of,  267. 
Birkbeck,   Mr.,   on   the   finding  of 
new  mates  by  golden  eagles,  403. 
Birthplace  of  man,  157. 
Births,    numerical   proportions    of 
the    sexes    in,    in    animals     and 
man,  210,  211;     male  and  female, 
numerical      proportion      of,      in 
England,   237. 
Bischoff,  Prof.,  on  the  agreement 
between  the  brains  of  man  and 
of   the    orang,   6;     figure   of   the 
embryo   of  the  dog,   10;     on   the 
convolutions  of  the  brain  in  the 
human  foetus,  11;    on  the  differ- 
ence between  the  skulls  of  man 
and    the    quadrumana,    145;     re- 
semblance    between    the     ape's 
and  man's,  194. 
Bishop,  J.,  on  the  vocal  organs  of 
frogs,   346;     on  the  vocal  organs 
of    corvine   birds,     366;     on     the 
trachea  of  the  Merganser,  369. 
Bison,  American,  co-operation  of, 

98;    mane  of  the  male,  517. 
Bitterns,  dwarf,  coloration  of  the 

sexes  of,  454. 
Biziura  lobata,  musky  odor  of  the 
male,    355;     large    size    of    male, 
358. 
Blackbird,    sexual     differences    in 
the,  214;    proportion  of  the  sexes 
in  the,  243;    acquisition  of  a  song 
by,  366;    color  of  the  beak  in  the 
sexes    of    the,    377,    487;     pairing 
with    a  thrush,    409;     colors    and 
nidification  of  the,  449;  young  of 
the,  487;  sexual  difference  in  col- 
oration of  the,   487. 
Black-buck,  Indian,  sexual  differ- 
ence in  the  color  of  the,  531. 
Blackcap,  arrival  of  the  male,  be- 
fore the  female,   208;    young  of 
the,  482. 
Black-cock,  polygamous,  215;    pro- 
portion of  the  sexes  in  the,  243; 
pugnacity  and  love-dance  of  the, 
359;    call  of  the,  370;   moulting  of 
the,  386;     duration  of  the  court- 
ship  of  the,   400;     and  pheasant, 
hybrids  of,  409;    sexual  difference 
in  coloration  of  the,  491;  crimson 
eye-cere  of   the,  487. 
Blacklock,  Dr.,  on  music,  567. 
Black-grouse,         characters        of 

young,  459,  465. 
Blackwall,  J.,  on  the  speaking  of 
the  magpie,  87;  on  the  desertion 
of  their  young  by  swallows,  104; 
on  the  superior  activity  of  male 
spiders,  217;  on  the  proportion 
of  the  sexes  in  spiders,  251;  on 
sexual  variation  of  color  in 
spiders,  269;  on  male  spiders, 
269. 


Bladder-nose    Seal,    hood    of    the, 

523. 
Blaine,  on  the  affections  of  dogs, 

519. 
Blair,  Dr.,  on  the  relative  liability 
of    Europeans   to    yellow    fever, 
189. 
Blake,  C.  C,  on  the  jaw  from  La 

Naulette,  39. 
Blakiston,  Capt.,  on  the  American 
snipe,    373;      on    the    dances     of 
Tetrao  phasianellus,  376. 
Blasius,    Dr.,    on    the     species     of 

European  birds,   416. 
Bledius  taurus,  hornlike  processes 

of   male,   296. 
Bleeding,  tendency  to  profuse,  232. 
BJenkiron,   Mr.,  on  sexual  prefer- 
ence in  horses,  520. 
Blennies,    crest   developed   on   the 
head  of  males,  during  the  breed- 
ing season,  336. 
Blethisa   multipunctata,    stridula- 

tion  of,  300. 
Bloch,    on   the  proportions   of  the 

sexes  in  fishes,  244. 
Blood,  arterial,  red  color  of,  258. 

pheasant,   number  of   spurs 

in,  360. 
Bluebreast,    red-throated,     sexual 

differences   of   the,   466. 
Blumenbach,    on  Man,   27;    on  the 
large  size  of  the   nasal   cavities 
in  American   aborigines,   33;     on 
the  position  of  man,  145;    on  the 
number  of  species  of  man,  170. 
Blyth,  E.,  on  the  structure  of  the 
hand    in    the    species    of    Hylo- 
bates,  49;  observations  on  Indian 
crows,  99;  on  the  development  of 
the    horns    in    the    Koodoo    and 
Eland  antelopes,  229;  on  the  pug- 
nacity of  the  males  of  Gallinula 
cristata,  357;    on  the  presence  of 
spurs  in  the  female  Euplocamus 
erythrophthalmus,    360;    on     the 
pugnacity  of  the  amadavat,  362; 
on   the    spoonbill,     370;     on     the 
moulting  of  Anthus,  386;    on  the 
moulting    of    bustards,     plovers, 
and  Gallus  bankiva,  387;    on  the 
Indian   honey-buzzard,     418;     on 
sexual  differences  in  the  color  of 
the  eyes    of    hornbills,     420;    on 
Oriolus  melanocephalus,  454;    on 
Palseornis  javanicus,  455;    on  the 
genus  Ardetta,  455;    on  the  pere- 
grine falcon,   455;     on  young  fe- 
male birds  acquiring  male  char- 
acters,   455;      on    the    immature 
plumage  of  birds,  459;    on  repre- 
sentative   species    of    birds,   462; 
on  the  young  of  Turnix,  470;    on 
anomalous     young     of     Lanius 
rufus   and    Colymbus     glacialis, 
477;    on  the  sexes  and  young  of 
the    sparrows,     477;     on     dimor- 
phism  in  some   herons,   479;     on 
the  ascertainment  of  the  sex  of 
nestling    bullfinches   by    pulling 


622 


INDEX. 


out  breast-feathers,  478;  on  ori- 
oles breeding-  in  immature  plum- 
age, 479;  on  the  sexes  and  young 
of  Buphus  and  Anastomus,  481; 
on  the  young-  of  the  blackcap 
and  blackbird,  482;  on  the  young 
of  the  stonechat,  4S7;  on  the 
v/hite  plumage  of  Anastomus, 
4oS;  on  the  horns  of  Bovine  ani- 
mals, 501;  on  the  horns  of  An- 
tilope  bezoartica,  503;  on  the 
mode  of  fighting  of  Ovis  cyclo- 
ceros,  504;  on  the  voice  of  the 
Gibbons,  522;  on  the  crest  of  the 
male  wild  goat,  526;  on  the  colors 
of  Portax  picta,  530;  on  the  col- 
ors of  Antilope  bezoartica,  531; 
on  the  color  of  the  Axis  deer, 
532;  on  sexual  difference  of  color 
in  Hylobates  hoolock,  532;  on  the 
hog--deer,  541;  on  the  beard  and 
whiskers  in  a  monkey  becoming 
white  with  age,  554. 

Boar,  wild,  polygamous  in  India, 
213;  use  of  the  tusks  by  the,  509; 
fighting  of,   514. 

Boardman,  Mr.,  Albino  birds  in  U. 
S.,  413. 

Boitard  and  Corbie,  MM.,  on  the 
transmission  of  sexual  peculiar- 
ities in  pigeons,  225;  on  the  an- 
tipathy shov/n  by  some  female 
pigeons   to   certain   males,    412. 

Bold,  Mr.,  on  the  singing-  of  a 
sterile  hybrid  canary,  365. 

Bcmbet,  on  the  variability  of  the 
standard  of  beauty  in  Europe, 
591. 

Bonibus,  difference  of  the  sexes 
in,  290. 

Bombycid^,  coloration  of,  310; 
pairing  of  the,  314;  colors  of,  315. 

Bombycilla  carolinensis,  red  ap- 
pendages of,   455. 

Bombyx  cynthia,  276;  proportion 
of  the  sexes  in,  245,  249;  pairing 
of,  314.  ^      . 

mori,    difference    of    size   of 

the  male  and  female  cocoons  of, 
275;     pairing   of,    314. 

Pernyi,   proportion  of  sexes 


of,  249. 

Yama-mai,   276;   M.   Person- 


nat  on,  246;  proportion  of  sexes 
of,  249. 

Bonaparte,  C.  L..,  on  the  call- 
notes   of  the   wild  turkey,    370. 

Bond,  F.,  on  the  finding  of  new 
mates  by  crows,   402. 

Bone,  implements  of,  skill  dis- 
played in  making,  48. 

Boner,  C,  on  the  transfer  of  male 
characters  to  an  old  female 
chamois,  500;  on  the  habits  of 
stags,  511;  on  the  pairing  of  red 
deer,  518. 

Bones,  increase  of,  in  length  and 
thickness,  when  carrying  a 
greater  weight,  31. 

Bonizzi,  P.,  difference  of  color  in 
sexes    of    pigeons,    225. 


Bonnet  monkey,   146. 

Bonwick,  J.,  extinction  of  Tasma- 
nians,    179,   180. 

Boomerang,   141. 

Boreus  hyemalis,  scarcity  of  the 
male,    251. 

Bory  St.  Vincent,  on  the  number 
of  species  of  man,  170;  on  the 
colors  of  Labrus  pavo,    339. 

Bos  etruscus,  501. 

gaurus,  horns  of,  501. 

■ •  moschatus,    525. 

• primig-enius,  497. 

sondaicus,      horns     of,     501; 

colors  of,  531. 

Botocudos,  140;  mode  of  life  of, 
192;  disfigurement  of  the  ears 
and  lower  lip  of  the,  570. 

Boucher  de  Perthes,  J.  C,  de,  on 
the  antiquity  of  man,  2. 

Bourbon,  proportion  of  the  sexes 
in  one  species  of  Papilio  from, 
246. 

Bourien,  on  the  marriage-customs 
of  the  savages  of  the  Malay 
Archipelago,   593. 

Bovidse,   dewlaps  of,   527. 

Bower-birds,  401;  habits  of  the, 
377;  ornamented  playing-places 
of,    89,   408. 

Bows,   use  of,  174. 

Brachycephalic  structure,  pos- 
sible explanation  of,  54. 

Brachyura,  265. 

Brachyurus  calvus,  scarlet  face 
of,   546. 

Bradley,  Mr.,  abductor  ossis  met- 
atarsi quinti  in  man,  41. 

Brain,  of  man,  agreement  of  the, 
with  that  of  lower  animals,  6; 
convolutions  of,  in  the  human 
foetus,  11;  influence  of  develop- 
ment of  mental  faculties  upon 
the  size  of  the,  52;  influence  of 
the  development  of,  on  the  spi- 
nal column  and  skull,  53;  larger 
in  some  existing  mammals  than 
in  their  tertiary  prototypes,  78; 
relation  of  the  development  of 
the,  to  the  prog-ress  of  language, 
85;  disease  of  the,  affecting 
speech,  86;  difference  in  the  con- 
volutions of,  in  different  races 
of  men,  163;  supplement  on,  by 
Prof.  Huxley,  194;  development 
of  the  g-yri  and  sulci,  200. 

Brakenridge,  Dr.,  on  the  influence 
of  climate,   31. 

Brandt,  A.,  on  hairy  men,  19. 

Eraubach,  Prof.,  on  the  quasi-re- 
ligious feeling  of  a  dog  towards 
his  master,  93;  on  the  self-res- 
traint of  dog-s,  100. 

Brauer,  P.,  on  dimorphism  m 
Neurothemis,   288. 

Brazil,  skulls  found  in  caves  of, 
165;  population  of,  169;  compres- 
sion of  the  nose  by  the  natives 
of,    577. 

Break  between  man  and  the  apes, 
152. 


INDEX. 


623 


Bream,  proportion  of  the  sexes  in 
the,  245. 

Breeding,   age   of,    in   birds,   478. 

season,     sexual     characters 

making  their  appearance  in  the, 
in  birds,  385. 

Brehni,  on  the  effects  of  intoxicat- 
ing Kquors  on  monkeys,  7;  on 
the  recognition  of  women  by- 
male  Cynocephali,  8;  on  the  di- 
versity of  the  mental  faculties 
of  monkeys,  26;  on  the  habits  of 
baboons,  49;  on  revenge  taken 
by  monkeys,  67;  on  manifesta- 
'tions  of  maternal  afCection  by 
monkeys  and  baboons,  68;  on  tlie 
instinctive  dread  of  monkeys  for 
serpents,  69;  on  the  use  of  stones 
as  missiles  by  baboons,  79;  on  a 
baboon  using  a  mat  for  shelter 
from  the  sun,  80;  on  the  signal- 
cries  of  monkeys,  85;  on  sentin- 
els posted  by  monkeys,  98;  on  co- 
operation of  animals,  98;  on  an 
eagle  attacking  a  young  Cercop- 
ithecus,  98;  on  baboons  in  con- 
finement protecting  one  of  their 
number  from  punishment,  100; 
on  the  habits  of  baboons  when 
plundering,  101;  on  polygamy  in 
Cynocephalus  and  Cebus,  213;  on 
the  numerical  proportion  of  the 
sexes  in  birds,  242;  on  the  love- 
dance  of  the  black-cock,  259;  on 
Palamedea  cornuta,  360;  on  the 
habits  of  the  Black-grouse,  362; 
on  sounds  produced  by  birds  of 
paradise,  371;  on  assemblages  of 
grouse,  400;  on  the  finding  of 
new  mates  by  birds,  403;  on  the 
fighting  of  wild  boars,  514;  on 
the  habits  of  Cynocephalus 
hamadryas,  585. 

Brent,  Mr.,  on  the  courtship  of 
fowls,  411. 

Breslau,  numerical  proportion  of 
male  and  female  births  in,  237. 

Bridgman,  Laura,  85. 

Brimstone  butterfly,  319;  sexual 
difference  of  color  in  the,   319. 

British,  ancient,  tattooing  prac- 
ticed by,   569. 

Broca,  Prof.,  on  the  occurrence  of 
the  supra-condyloid  foramen  in 
the  hunian  humerus,  21;  anthro- 
pomorphous apes  more  bipedal 
than  Quadrupedal,  51;  on  the 
capacity  of  Parisian  skulls  at 
different  periods,  53;  comparison 
of  modern  and  mediaeval  skulls, 
53;  on  tails  of  quadrupeds,  56; 
on  the  influence  of  natural  se- 
lection, 58;  on  hybridity  in  man, 
166;  on  human  remains  from  Les 
Eyzies,  177;  on  the  cause  of  the 
difference  between  Europeans 
and  Hindoos,  187. 

Brodie,  Sir  B.,  on  the  origin  of 
the  moral  sense  in  man,  95. 

Bronn,  H.  G.,  on  the  copulation  of 
insects  of  distinct  species,  272. 


Bronze  period,  men  of,  in  Europe, 

125. 

Brown,  R.,  sentinels  of  seals  gen- 
erally females,  97;  on  the  battles 
of  seals,  496;  on  the  narwhal, 
498;  on  the  occasional  absence 
of  the  tusks  in  the  female  wal- 
rus, 498;  on  the  bladder-nose 
seal,  524;  on  the  colors  of  the 
sexes  in  Phoca  Grcenlandica, 
530;  on  the  appreciation  of  music 
by  seals,  563;  on  plants  used  as 
love-philters,  by  North  Ameri- 
can women,  572. 

,  Dr.  Crichton,  injury  to  in- 
fants during   parturition,   238. 

Brown-Sequard,  Dr.,  on  the  inher- 
itance of  the  effects  of  opera- 
tions by  guinea-pigs,  57,  597. 

Bruce,  on  the  use  of  the  ele- 
phant's  tusks,   503. 

Brulerie,  P.  de  la,  on  the  habits  of 
Ateuchus  cicatricosus,  297;  on 
the  stridulation  of  Ateuchus, 
303. 

Brunnich,  on  the  pied  ravens  of 
the  Feroe  islands,  418. 

Bryant,  Dr.,  preference  of  tame 
pigeon  for  wild  mate,  413. 

,   Capt.,    on  the   courtship  of 

Callorhinus  ursinus,  518. 

Bubas  bison,  thoracic  projection 
of,  295. 

Bucephalus  capensis,  difference  of 
the  sexes  of,  in  color,  347. 

Buceros,  nidification  and  incuba- 
tion of,   448. 

bicornis,    sexual   differences 

in  the  coloring  of  the  casque, 
beak,  and  m.outh  in,  420. 

•  corrugatus,  sexual  differ- 
ences in  the  beak  of,  377. 

Buchner,  L.,  on  the  origin  ot 
man,  2;  on  the  use  of  the  human 
foot  as  a  prehensile  organ,  50; 
on  the  mode  of  progression  of 
the  apes,  50;  on  want  of  self- 
consciousness,  &c.,  in  savages, 
81. 

Bucholz,  Dr.,  quarrels  of  cham  • 
eeleons,  353. 

Buckland,  F.,  on  the  numerical 
proportion  of  the  sexes  in  rats, 
242;  on  the  proportion  of  the 
sexes  in  the  trout,  244;  on 
Chimsera   monstrosa,    336. 

,    W.,    on   the    complexity   of 

crinoids,  89. 

Buckler,  W.,  proportion  of  sexes 
of  Lepidoptera  reared  by,  249. 

Buckinghamshire,  numerical  pro- 
portion of  male  and  female 
births  in,  237. 

Bucorax  abyssinicus,  inflation  of 
the  neck-wattle  of  the  male  dur- 
ing courtship,  377. 

Budytes  Rail,  208. 

Buffalo,   Cape,  504. 

,   Indian,   horns  of   the,   501. 

,  Italian,  mode  of  fighting  of 

the,  504. 


624 


INDEX. 


Buffon,  on  the  number  of  species 

of  man,  170, 
Bufo  sikimmensis,  345. 
Bugs,   278. 

Buist,  R.,  on  the  proportion  of 
the  sexes  in  salmon,  244;  on  the 
pugnacity  of  the  male  salmon, 
329. 
Bulbul,  pugnacity  of  the  male, 
357;  display  of  under  tail-coverts 
by  the  male,  396. 
Bull,  mode  of  fighting  of  the,  504; 

curled  frontal  hair  of  the,  526. 
Buller,  Dr.,  on  the  Huia,  204;  the 

attachment  of  birds,  405. 
Bullfinch,     sexual     differences   in 
the,  214;  piping,  364;  female,  sing- 
ing of  the,  365;  courtship  of  the, 
395;    widowed,     finding     a     new 
mate,     403;     attacking    a    reed- 
bunting,  407;  nestling,  sex  ascer- 
tained   by    pulling     out    breast- 
feathers,  478. 
Bullfinches       distinguishing     per- 
sons, 406;   rivalry  of  female,  414. 
Bulls,    two   young,    attacking    an 
old  one,  98;  wild,  battles  of,  497. 
Bull-trout,  male,  coloring  of,  dur- 
ing   the   breeding    season,    336. 
Bunting,   reed,   head     feathers   of 
the  male,  396;  attacked  by  a  bull- 
finch,  407. 
Buntings,     characters    of  young, 

459. 
Buphus  coromandus,     sexes     and 
young  of,  481;  change  of  color  in, 
489,  490. 
Burchell,    Dr.,   on   the   zebra,   540; 
on  the  extravagance  of  a  bush- 
woman  in  adorning  herself,  572; 
celibacy    unknown    among    the 
savages  of  South  Africa,  588;  on 
the     marriage-customs     of     the 
Bushwomen,  393. 
Burke,  on  the  number  of  species 

of  man,  170. 
Burmese,   color   of  the   beard  in, 

554. 
Burton,  Capt.,   on  negro  ideas  of 
female  beauty,  574;  on  a  univer- 
sal ideal  of  beauty,  576. 
Bushmen,  61. 

Bushwoman,    extravagant     orna- 
mentation of  a,  572. 
Bushwomen,    hair    of,    163;     mar- 
riage-customs of,  593. 
Bustard,       throat-pouch     of     the 
male,    368;    humming   noise  pro- 
duced   by    a    male,    373;    Indiail, 
ear-tufts  of,   379. 
Bustards,    occurrence    of    sexual 
differences     and     of     polygamy 
among  the,  215;  love-gestures  of 
the  male,   376;   double  moult  in, 
385,  386. 
Butler,    A.    G.,    on    sexual    differ- 
ences  in  the   wings   of  Aricoris 
epitus,  277;   courtship  of  butter- 
flies, 304;  on  the  coloring  of  the 
sexes  in  species  of  Thecla,  307; 


on  the  resemblance  of  Iphias 
glaucippe  to  a  leaf,  309;  on  the 
rejection  of  certain  moths  and 
caterpillars  by  lizards  and  frogs, 
322. 

Butterfly,  noise  produced  by  a, 
304;  Emperor,  304,  305;  meadow 
brown,  instability  of  the  ocel- 
lated  spots  of,   422. 

Butterflies,  proportion  of  the 
sexes  in,  246;  forelegs  atrophied 
in  some  male,  274;  sexual  dif- 
ference in  the  neuration  of  the 
wings  of,  274;  pugnacity  of  male, 
304;  protective  resemblances  of 
the  lower  surface  of,  308;  display 
of  the  wings  by,  311;  white, 
alighting  upon  bits  of  paper, 
314;  attracted  by  a  dead  speci- 
men of  the  same  species,  314; 
courtship  of,  314;  male  and  fe- 
male, inhabiting  different  sta- 
tions, 317. 

Buxton,  C,  observations  on  ma- 
caws, 99;  on  an  instance  of  be- 
nevolence in  a  parrot,  406. 

Buzzard,  Indian  honey-,  variation 
in  the  crest  of,  418. 


Cabbage  butterflies,  309. 

Cachalot,  large  head  of  the  male, 
498. 

Cadences,  musical,  perception  of, 
by  animals,  564. 

Csecum,  20;  large,  in  the  early 
progenitors  of  man,  156. 

Cairina  moschata,  pugnacity  of 
the  male,   358. 

Californian  Indians,  decrease  of, 
254. 

Callianassa,  chelae  of,  flgured,  263. 

Callidryas,   colors  of   sexes,   315. 

Callionymus  lyra,  characters  of 
the  male,  332. 

Callorhinus  ursinus,  relative  size 
of  the  sexes  of,  511;  courtship  of, 
518. 

Calotes  maria,  353. 

nigrilabris,  sexual  differ- 
ence in  the  color  of,  353. 

Cambridge,  O.  Pickard,  on  the 
sexes  of  spiders,  251;  on  the  size 
of  male  Nephila,  270. 

Camel,  canine  teeth  of  male,  498, 
509. 

Campbell,  J.,  on  the  Indian  ele- 
phant, 213;  on  the  proportion  of 
male  and  female  births  in  the 
harems  of  Siam,  239. 

Campylopterus   hemileucurus,  243. 

Canaries  distinguishing  persons, 
406. 

Canary,  polygamy  of  the,  215; 
change  of  plumage  in,  after 
moulting,  233;  female,  selecting 
the  best  singing  male,  364;  ster- 
ile hybrid,  singing  of  a,  365;  fe- 


INDEX. 


625 


male,  singing  of  the,  365;  select- 
ing a  greenfinch,  410;  and  siskin, 
pairing  of,  410. 

Canestrini,  G.,  on  rudimentary 
characters  and  the  origin  of 
man,  2;  on  rudimentary  charac- 
ters, 11;  on  the  movement  of  the 
ear  in  man,  14;  on  the  variabil- 
ity of  the  vermiform  appendage 
in  man,  20;  on  the  abnormal  di- 
vision of  the  malar  bone  in  man, 
37;  on  abnormal  conditions  of  the 
human  uterus,  38;  on  the  per- 
sistence of  the  frontal  suture  in 
man,  38;  on  the  proportion  of  the 
sexes  in  silk-moths,  245,  247; 
secondary  sexual  characters  of 
spiders,  269. 

Cancer  pagurus,  263. 

Canfield,  Dr.,  on  the  horns  of  the 
Antilocapra,  229. 

Canine  teeth  in  man,  39;  diminu- 
tion of,  in  man,  51;  diminution 
of,  in  horses,  52;  disappearance 
of,  in  male  ruminants,  52;  large, 
in  the  early  progenitors  of  man, 
156. 

Canines,  and  horns,  inverse  devel- 
opment of,  509. 

Canoes,  use  of,  47,  175. 

Cantharis,  difference  of  color  in 
the  sexes  of  a  species  of,  291. 

Cantharus  lineatus,  337. 

Capercailzie,  polygamous,  215; 
proportion  of  the  sexes  in  the, 
243;  pugnacity  of  the  male,  359; 
pairing  of  the,  362;  autumn 
meetings  of  the,  366;  call  of  the, 
370;  duration  of  the  courtship  of, 
400;  behavior  of  the  female, 
414j.  inconvenience  of  black  col- 
or "to  the  female,  438;  sexual  dif- 
ference in  the  coloration  of  the, 
487;  crimson  eye-cere  of  the 
male,  487. 

Capitonidae,  colors  and  nidifica- 
tion  of  the,  449. 

Capra  £egagrus,  503;  crest  of  the 
male,  526;  sexual  difference  in 
the  color   of,   531. 

Capreolus  Sibricus  subecaudatus, 
537. 

Caprice,  common  to  man  and  ani- 
mals, 91. 

Caprimulgus,  noise  made  by  the 
males  of  some  species  of,  with 
their  wings,  371. 

virginianus,  pairing  of,  362. 

CarabidEB,  300. 

Carbonnier,  on  the  natural  his- 
tory of  the  pike,  244;  on  the  rel- 
ative size  of  the  sexes  in  fishes, 
331;  courtship  of  Chinese  Ma- 
cropus,   337. 

Carcineutes,  sexual  difference  of 
color  in,  451. 

Carcinus  meenas,  265. 

Cardinalis  virginianus,  220. 

Carduelis  elegans,  sexual  differ- 
ences of  the  beak  in,  356. 

40 


Carnivora,     marine,     polygamous 

habits  of,  214;  sexual  differences 

in  the  colors  of,  530. 
Car"p,  numerical  proportion  of  the 

sexes  in  the,  245. 
Carr,  R.,  on  the  peewit,  362. 
Carrier  pigeon,    late   development 

of  the  wattle  in  the,  232. 
Carrion    beetles,    stridulation    of, 

299. 

Carus,  Prof.  V.,  on  the  develop- 
ment of  the  horns  in  merino 
sheep,  230. 

Cassowary,  sexes  and  incubation 
of  the,  472. 

Castina,  mode  of  holding  wings, 
311. 

Castoreum,  524. 

Casuarius  galeatus,  472. 

Cat,  convoluted  body  in  the  ex- 
tremity of  the  tail  of  a,  22; 
sick,  sym.pathj'-  of  a  dog  with  a, 
100. 

Cataract  in  Cebus  Azarse,  7. 

Catarrh,  liability  of  Cebus  Azarae 
to,  7. 

Catarrhine  monkeys,  149. 

Caterpillars,  bright  colors  of,  321. 

Cathartes  aura,  410. 

jota,     love-gestures   of   the 

male,  375. 

Catlin,  G.,  correlation  of  color  and 
texture  of  hair  in  the  Mandans, 
192;  on  the  development  of  the 
beard  among  North  American 
Indians,  555;  on  the  great  length 
of  the  hair  in  some  North  Amer- 
ican tribes,  575. 

Caton,  J.  D.,  on  the  development 
of  the  horns  in  Cervus  virgin- 
ianus and  strongyloceros,  229;  on 
the  presence  of  traces  of  horns 
in  the  female  wapiti,  500;  on  the 
fighting  of  deer,  506;  on  the  crest 
of  the  male  wapiti,  526;  on  the 
colors  of  the  "Virginian  deer,  530; 
on  sexual  differences  of  color 
in  the  wapiti,  531;  on  the  spots 
of  the  Virginian  deer,  541, 

Cats,  dreaming,  72;  tortoise-shell, 
225,  227,  232;  enticed  by  valerian, 
530;  colors  of,  538. 

Cattle,  rapid  increase  of,  in  South 
America,  45;  domestic,  lighter  in 
winter  in  Siberia,  224;  horns  of, 
229,  501;  domestic,  sexual  differ- 
ences of,  late  developed,  232; 
numerical  proportion  of  the 
sexes  in,  241. 

Caudal  vertebrae,  number  of,  in 
macaques  and  baboons,  56;  bas- 
al, of  monkeys,  imbedded  in  the 
body,   57. 

Cavolini,  observations  on  Serran- 
us,  157. 

Cebus,  maternal  affection  in  a,  68; 
gradation  of  species  of,  171. 

Apella,  201. 

Azarae,    liability  of,    to   the 

same  diseases   as  man,    7;   dis- 


626 


INDEX. 


timet  sounds  produced  hy,  82; 
early  maturity  of  the  female, 
553. 
Cebus  capucinus  polygamous,  213; 
sexual  differences  of  color  in, 
532;   hair   on    the   head    of,    544. 

vellerosus,  hair  on  the  head 

of,  544. 

Cecidomyidse,  proportions  of  the 
sexes  in,  250. 

Celibacy,  unknown  among-  the 
savages  of  South  Africa  and 
South  America,  588. 

Centipedes,  270. 

Cephalopoda,  absence  of  second- 
ary sexual  characters   in,    260. 

Cephalopterus  ornatus,  368. 

penduliger,  368. 

Cerambyx  heros,  stridulant  organ 
of,  300. 

Ceratodus,  paddle  of,  35. 

Ceratophora  aspera,  nasal  ap- 
pendages of,  351. 

Stoddartii,     nasal   horn  of, 

351. 

Cerceris,  habits  of,  289. 
Cercocebus      sethiops,      whiskers, 

&c.,   of,  546. 
Cercopithecus,     young,    seized  by 

an    eagle    and   rescued    by     the 

troop,  98;  definition  of  species  of, 

171. 

cephus,  sexual  difference  of 

color  in,  532,  547. 

cynosurus  and  griseoviridis, 


color  of  the  scrotum  in,   533. 

Diana,  sexual  differences  of 


color  in,  532,  547. 

griseo-viridis,  98. 

petaurista,  whiskers,  &c.,  of, 


545. 


Ceres,   of  birds,  bright  colors  of, 

487. 
Ceriornis  Temminckii,  swelling  of 

the  wattles  of  the  male  during 

courtship,  377. 
Cervulus,  weapons  of,  509. 

m.oschatus,      rudimentary 

horns  of  the  fem.ale,  500. 

Cervus   alces,  229. 

campestris,   odor  of,   524. 

canadensis,  traces  of  horns 

in  the  female,  500;  attacking  a 
man,  506;  sexual  difference  in  the 
color  of,  531. 

elaphus,     battles     of   male. 


497;     horns   of,   with     numerous 
points,  506. 

Eldi,  229. 

mantchuricus,  541. 

paludosus,  colors  of,  532. 

strongyloceros,  229. 

virginianus,  229;  horns  of,  in 


course  of  modification,  508. 
Ceryle,  male  black-belted  in  some 

species  of,  451. 
Cetacea,  nakedness  of,   54. 
Ceylon,  frequent  absence  of  beard 

in  the  natives  of,  555. 
Chaffinch,       proportion      of      the 


sexes  in  the,  224;  courtship  of 
the,   395. 

Chaffinches,  365;  new  mates 
found  by,  403. 

Chalcophaps  indicus,  characters 
of  young,  459. 

Chalcosoma  atlas,  sexual  differ- 
ences of,  293. 

Chameeleo,  sexual  differences  in 
the  genus,  352. 

bifurcus,  352. 

Owenii,  352. 

pumilus,  353. 

Chamsepetes      unicolor,      modified 

wingfeather  in  the  male,   373. 

Chameleons,   350. 

Chamois,  danger-signals  of,  97; 
transfer  of  male  characters  to 
an  old  female,  499,  500. 

Champneys,  Mr.,  acromio-basilar 
muscle  and  quadrupedal  gait,  40. 

Chapuis,  Dr.,  on  the  transmission 
of  sexual  peculiarities  in  pig- 
eons, 225;  on  streaked  Belgian 
pigeons,   233,   440. 

Char,  male,  coloring  of,  during 
the  breeding  sea.son,   336. 

Characters,  male,  developed  in 
females,  223;  secondary  sexual, 
transmitted  through  both  sexes, 
222;  natural,  artificial,  exagger- 
ation of,  by  man,  577. 

Charadrius  hiaticula  and  pluvial- 
is,   sexes  and  young  of,  480. 

Chardin  on  the  Persians,  581. 

Charms,  worn  by  women,   572. 

Charruas,  freedom  of  divorce 
among  the.  592. 

Chasmorhynchus,  difference  of 
color  in  the  sexes  of,  384;  colors 
of,  488. 

niveus,  384. 

tricarunculatus,  384. 

Chastity,  early  estimation  of,  115. 

Chatterers,  sexual  differences  in, 
214. 

Cheever,  Rev.  H.  T.,  census  of  the 
Sandwich  Islands,  254. 

Cheiroptera,  absence  of  secondary 
sexual  characters  in,  214. 

Chelae  of  Crustacea,  263,  268. 

Chelonia,  sexual  differences  in, 
346. 

Chenalopex  segyptiacus,  wing- 
knobs  of,  360. 

Chera  progne,  387,  413. 

Chest,  proportions  of,  in  soldiers 
and  sailors,  31;  large,  of  the 
Quechua  and  Aymara  Indians, 
33. 

Chevrotains,   canine  teeth  of,  509. 

Chiasognathus,  stridulation  of, 
303. 

Grantii,   mandibles  of,  298. 

Children,  legitimate  and  illegiti- 
mate, proportion  of  the  sexes  in, 
237. 

Chiloe,  lice  of  the  natives  of,  166; 
population  of,  169. 

Chimsera  monstrosa,  bony  proc- 
ess on  the  head  of  the  male,  334. 


INDEX. 


627 


Chimseroid  fishes,  prehensile  or- 
gans of  male,  327. 

Chimpanzee,  556;  ears  of  the,  14; 
representatives  of  the  eyebrows 
in  the,  18;  hands  of  the,  48;  ab- 
sence of  mastoid  processes  in 
the,  51;  platforms  buiit  by  tlie, 
64;  cracliing  nuts  with  a  stone, 
78;  direction  of  the  hair  on  the 
arms  of  the,  147;  supposed  evo- 
lution of  the,  173;  polygamous 
and  social   habits   of    the,    585. 

China,  North,  idea  of  female 
beauty  in,  572. 

,    Southern,     inhabitants    of, 

192. 

Chinese,  use  of  flint  tools  by  the, 
141;  difficulty  of  distinguishing 
the  races  of  the,  163;  color  of  the 
beard  in,  554;  general  beardless- 
ness  of  the,  555;  opinions  of  the, 
on  the  appearance  of  Europeans 
and  Cingalese,  572;  compression 
of  the  feet  of,  577. 

Chinsurdi,  his  opinion  of  beards, 
570,    575. 

Chlamydera  maculata,  377. 

Chloeon,  pedunculated  eyes  of  the 
male   of,   272. 

Chlcephaga,  coloration  of  the 
sexes  in,  454. 

Chlorocaelus  Tanana  (figured), 
283. 

Chorda  Dorsalis,  156. 

Chough,  red  beak  of  the,  487. 

Chromidse,  frontal  protuberance 
in  male,  336;  sexual  differences 
in  color  of,  342. 

Chrysemys  picta,  long  claws  of 
the  male,  346. 

Chrysococcyx,  characters  of 
young  of,    459. 

Chrysomelidas,  stridulation  of, 
299. 

Cicada  pruinosa,  279. 

septendecim,  279. 

CicadEe,  songs  of  the,  279;  rudi- 
mentary sound-organs  in  fe- 
males of,  285. 

Cicatrix  of  a  burn,  causing  modi- 
fication  of  the  facial  bones,   53. 

Cichla,  frontal  protuberance  of 
male,  336. 

Cimetiere  du  Sud,  Paris,  21. 

Cincloramphus  cruralis,  large  size 
of  male,  358. 

Cinclus  aquaticus,  449. 

Cingalese,  Chinese  opinion  of  the 
appearance  of  the,  572. 

Cirripedes,  complemental  males 
of,   204. 

Civilization,  effects  of,  upon  nat- 
ural selection,  130;  infiuence  of, 
in  the  competition  of  nations, 
178. 

Clanging  of  geese,   &c.,  364. 

Claparede,  E.,  on  natural  selec- 
tion applied  to  man,  47. 

Clarke,  on  the  marriage-customs 
of  the  Kalmucks,  593. 


Classification,  144. 

Claus,  C,  on  the  sexes  of  Sap- 
hirina,   268. 

Cleft-palate,   inherited,   34. 

Climacteris  erj^throps,  sexes  of, 
473. 

Climate,  30;  cool,  favorable  to  hu- 
man progress,  129;  power  of  sup- 
porting extremes  of,  by  man, 
177;  want  of  connection  of,  with 
color,  187. 

Cloaca,  existence  of  a,  in  the 
early  progenitors  of  man,  156. 

Cloacal  passage  exising  in  the  hu- 
man embryo,  9. 

Clubs,  used  as  weapons  before 
dispersion  of  mankind,  175. 

Clucking  of  fowls,  363. 

Clythra  4-punctata,  stridulation 
of,   299. 

Coan,  Mr.,  Sandwich-islanders, 
182. 

Cobbe,  Miss,  on  morality  in  hypo- 
thetical bee-community,  96. 

Cobra,  ingenuity  of  a,  348. 

Coccus,   143. 

Coccyx,  22;  in  the  human  em- 
bryo, 9;  convoluted  body  at  the 
extremity  of  the,  22;  imbedded 
in  the  body,  57. 

Cochin-China,  notions  of  beauty 
of  the  inhabitants  of,  572,  574. 

Cock,  blind,  fed  by  its  compan- 
ions, 100;  game,  killing  a  kite, 
359;  comb  and  wattles  of  the, 
398;  preference  shown  by  the,  for 
young  hens,  414;  game,  transpar- 
ent zone  in  the  hackles  of  a, 
424. 

Cock  of  the  rock,  400. 

Cockatoos,  487,  488,  489;  nestling, 
406;  black,  immature  plumage  of, 
461. 

Coelenterata,  absence  of  second- 
ary  sexual   characters   in,   257. 

Coffee,  fondness  of  monkeys  for, 
7. 

Cold,  supposed  effects  of,  31; 
power  of  supporting,  by  man, 
177. 

Coleoptera,  291;  stridulation  of, 
282;  stridulant  organs  of,  dis- 
cussed, 301. 

Colias  edusa  and  hyale,  315. 

Collingwood,  C,  on  the  pugnac- 
ity of  the  butterflies  of  Borneo, 
304;  on  butterflies  being  attract- 
ed by  a  dead  specimen  of  the 
same  species,  314. 

Colobus,  absence  of  the  thumb, 
49. 

Colombia,  flattened  heads  of  sav- 
ages of,  569. 

Colonists,  success  of  the  English, 
as,  138. 

Coloration,  prdtective,  in  birds, 
484. 

Color,  supposed  to  be  dependent 
on  light  and  heat,  31;  corx^elation 
of,  with  immunity  from  certain 


628 


INDEX. 


poisons  and  parasites,  188;  pur- 
pose of,  in  lepidoptera,  313;  rela- 
tion of,  to  sexual  functions,  in 
fishes,  339;  difference  of,  in  the 
sexes  of  snakes,  347;  sexual  dif- 
ferences of,  in  lizards,  353;  in- 
fluence of,  in  the  pairing  of 
birds  of  different  species,  410;  re- 
lation of,  to  nidification,  447,  551; 
sexual  differences  of,  in  mam- 
mals, 549;  recognition  of,  by 
quadrupeds,  535;  of  children,  in 
different  races  of  man,  552;  of 
the  skin  in  man,  598. 

Colors,  admired  alike  by  man  and 
animals,  90;  bright,  due  to  sex- 
ual selection,  258;  bright,  among 
the  lower  animals,  258,  259; 
bright,  protective  to  butterflies 
and  moths,  310;  bright,  in  male 
fishes,  332,  337;  transmission  of, 
in  birds,  442. 

Colquhoun,  example  of  reasoning 
in  a  retriever,  75. 

Columba  passerina,  young  of, 
461. 

Colymbus,  glacialis,  anomalous 
young  of,  477. 

Comb,  development  of,  in  fowls, 
233. 

Combs  and  wattles  in  male  birds, 
398. 

Community,  preservation  of  va- 
riations useful  to  the,  by  nat- 
ural selection,  60. 

Compositae,  gradation  of  species 
among  the,  170. 

Comte,  C,  on  the  expression  of 
the  ideal  of  beauty  by  sculpture, 
576. 

Conditions  of  life,  action  of 
changed,  upon  man,  29;  influ- 
ence of,  on  plumage  of  birds, 
467. 

Condor,  eyes  and  comb  of  the, 
472. 

Conjugations,  origin  of,   89. 

Conscience,  110,  122;  absence  of,  in 
some  criminals,   112. 

Constitution,  difference  of  in  dif- 
ferent races  of  men,  164. 

Consumption,  liability  of  Cebus 
Azaras  to,  7;  connection  be- 
tween complexion  and,  189. 

Convergence  of  characters,  173. 

Cooing  of  pigeons  and  doves,  370. 

Cook,  Capt.,  on  the  nobles  of  the 
Sandwich  Islands,  581. 

Cope,  B.  D.,  on  the  Dinosauria, 
154. 

Cophotis  ceylanica,  sexual  differ- 
ences of,  350,  353. 

Copris,  293. 

Isidis,  sexual  differences    of, 

294. 

lunaris,   stridulation  of,  300. 

Corals,   bright  colors   of,   257. 

Coral-snakes,  348. 

Cordylus,  sexual  difference  of  col- 
or in  a  species  of,  353. 


Corfu,  habits  of  the  Chaffinch  ifi, 

244. 
Cornelius,    on    the  proportions    of 

the    sexes    in    Lucanus    Cervus, 

249. 
Corpora      Wolffiana,     156;     agree- 
ment   of,    with    the    kidneys    of 

fishes,  11. 
Correlated  variation,  42. 
Correlation,    influence    of,    in    the 

production  of  races,  192. 
Corse,  on  the  mode  of  fighting  of 

the  elephant,  509. 
Corvus  corone,  402. 

graculus,   red  beak   of,   487. 

pica,    nuptial   assembly   of, 

401. 

Corydalis  cornutus,  large  jaws  of 

the  male,  273. 
Cosmetornis,  456. 

vexillarius,     elongation    of 

wing-feathers  in,  379,  397. 

Cotingidse,  sexual  differences  in, 
214;  coloration  of  the  sexes  of, 
453;  resemblance  of  the  females 
of  distinct  species  of,  464. 

Cottus  scorpius,  sexual  differences 
in,  334. 

Coulter,  Dr.,  on  the  Calif ornian 
Indians,  254. 

Counting,  origin  of,  140;  limited 
power  of,  in  primeval  man,  175. 

Courage,  variability  of,  in  the 
same  species,  67;  universal  high 
appreciation  of,  114;  importance 
of,  127;  characteristic  of  men, 
559. 

Courtship,  greater  eagerness  of 
males  in,  216;  of  fishes,  328,  337; 
of  birds,  363,  399. 

Cow,  winter  change  of  color,  537. 

Crab,    devil,    265. 

,  shore,  habits  of,  265. 

Crabro  cribrarius,  dilated  tibise 
of  the  male,  273. 

Crabs,  proportions  of  the  sexes  in, 
252. 

Cranz,  on  the  inheritance  of  dex- 
terity in  seal  catching,  32. 

Crawfurd,  on  the  number  of 
species  of  man,  170. 

Crenilabrus  massa  and  C.  melops, 
nests  built  by,  341. 

Crest,  origin  of,  in  Polish  fowls, 
226. 

Crests,  of  birds,  difference  of,  in 
the  sexes,  462;  dorsal  hairy,  of 
mammals,  526. 

Cricket,  field-,  stridulation  of  the, 
280;  pugnacity  of  male,  286. 

,   house-,   stridulation  of  the 

280,   282. 

Crickets,  sexual  differences  in, 
287. 

Crioceridae,  stridulation  of  the 
299. 

Crinoids,  complexity  of,  89. 

Croaking  of  frogs,  346. 

Crocodiles  musky  odor  of,  dur- 
ing  the   breeding   season,    347. 


INDEX. 


629 


Crocodilia,  347. 

Crossbills,     characters  of   young, 

459. 
Crosses  in  man,  169. 
Crossing  of  races,   effects  of  the, 

187. 
Crossoptilon  aurituni,  395,  446,  466; 
adornment  of  both  sexes  of,  230; 
sexes  alike  in,  454. 
Crotch,  G.  R.,  on  the  stridulation 
of  beetles,  299,  301;  on  the  strid- 
ulation of  Heliopathes,     302;   on 
the  stridulation  of  Acalles,   303; 
habit  of  female  deer  at  breeding 
time,  499. 
Crow  Indians,   long  hair  of    the, 
574. 

,  young  of  the,  475. 

Crows,   487;    vocal   organs   of  the, 
366;  living  in  triplets,  404. 

,   carrion,    new  mates  found 

by,  402. 

,  Indian,  feeding  their  blind 

companions,  99. 
Cruelty    of    savages     to   animals, 

114. 
Crustacea,  parasitic,  loss  of  limbs 
by    female,    204;    prehensile    feet 
and  antennse  of,  205;  male,  more 
active  than  female,  217;  parthen- 
ogenesis in,  251;  secondary  sex- 
ual characters  of,  262;  amphipod, 
males     sexually     mature     while 
young,   479;   auditory      hairs   of, 
563. 
Crystal  worn  in  the  lower  lip  by 
some   Central     African   women, 
570. 
Cuckoo  fowls,  233. 
Culicidee,    204,    278;     attracted     by 

each  other's  humming,  278. 
Cullen,   Dr.,   on  the   throat-pouch 

of  the  male  bustard,  368. 
Cultivation     of     plants,   probable 

origin  of,  130. 
Cupples,    Mr.,    on     the   numerical 
proportion  of  the  sexes  in  dogs, 
sheep,    and   cattle,     241;    on   the 
Scotch   deerhound,    512;    on   sex- 
ual preference  in  dogs,  520. 
Curculionidse,  sexual  difference  in 
length    of    snout    in    some,    204; 
hornlike  processes  in  male,  296; 
musical,  298,  299. 
Curiosity,    manifestations    of,    by 

animals,  69. 
Curlews,  double  moult  in,  385. 
Cursores,  comparative  absence  of 
sexual   differences   among     the, 
215. 
Curtis,    J.,    on    the   proportion    of 

the  sexes  in  Athalia,  250. 
Cuvier,  F.,  on  the  recognition  of 
women  by  male  quadrumana,  8. 
' ,  G.,  on  the  number  of  cau- 
dal vertebrae  in  the  mandrill, 
56;  on  instinct  and  intelligence, 
65;  views  of,  as  to  the  position 
of  man,  145;  on  the  position  of 
the  seals,  146;  on  Hectocotyle, 
260. 

41 


Cyanecula  suecica,  sexual  differ- 
ences of,  466. 

Cyanalcyon,  sexual  difference  in 
colors  of,  451;  immature  plu- 
mage of,  461. 

Cychrus,  sounds  produced  by,  301. 

Cycnia  mendica,  sexual  difference 
of,  in  color,  312. 

Cygnus  ferus,  trachea  of,  369. 

■ olor,   white  young  of,  477. 

Cyllo  Leda,  instability  of  the  ocel- 
lated  spots  of,  422. 

Cynanthus,  variation  in  the  gen- 
us, 417. 

Cynipidee,  proportions  of  the 
sexes  in,  250. 

Cynocephalus,  difference  of  the 
young  from  the  adult,  8;  male, 
recognition  of  women  by,  8; 
polygamous  habits  of  species  of, 
213. 

chacma,  68. 

gelada,  79. 

hamadryas,    79,    585;    sexual 

difference  of  color  in,  533. 

leucophaeus,     colors    of    the 


sexes  of,  534. 
mormon,  colors  of  the    male, 

534,  536,  545. 
porcarius,  mane  of  the  male, 

517. 
Cypridina,       proportions     of     the 

sexes  in,  251. 
Cyprinidaa,     proportion      of      the 

sexes  in  the,  245. 

,  Indian,  339. 

Cyprinodontidse,      sexual      differ- 
ences in  the,  332,  334. 
Cyprinus  auratus,  339. 
Cypris,  relations  of  the  sexes  in, 

251. 
Cyrtodactylus  rubidus,  350. 
Cystophora  cristata,  hood  of,  523. 


D 


Dacelo,  sexual  difference  of  color 

in,  451. 
Gaudichaudi,  young  male  of, 

462. 
Dal-ripa,    a   kind     of    ptarmigan, 

243. 
Damalis  albifrons,  peculiar  mark- 
ings of,  539. 
pygarga,  peculiar  markings 

of,  538. 
Dampness    of   climate,     supposed 

influence  of,  on  the  color  of  the 

skin,  31,  188. 
Danaidas,    305. 
Dances  of  birds,  377. 
Dancing,   universality  of,  174. 
Daniell,    Dr.,    his     experience     of 

residence  in  West  Africa,   190. 
Darfur,  protuberances  artificially 

produced  by  natives  of,  569. 
Darwin,  F.,  on  the  stridulation  of 

Dermestes  murinus,  299. 


630 


INDEX. 


Dasychira  pudibunda,  sexual  dif- 
ference of  color  in,  312. 

Davis,  A.  H,,  on  the  pugnacity  of 
the  male  stag-beetle,  297. 

,  J.   B.,   on  the     capacity     of 

the  skull  in  various  races  of 
men,  52;  on  the  beards  of  the 
Polynesians,  555. 

Death-rate  higher  in  towns  than 
in  rural  districts,  135. 

Death-tick.   303. 

De  Candolle,  Alph.,  on  a  case  of 
inherited  power  of  moving  the 
scalp,  13. 

Declensions,  origin  of,  89. 

Decoration  in  birds,  377. 

Decticus,   282. 

Deer,  228;  development  of  the 
horns  in,  228;  spots  of  young, 
458,  541;  horns  of,  499,  501;  use  of 
horns  of.  505,  513;  horns  of  a,  in 
course  of  modification,  508;  size 
of  the  horns  of,  511;  female,  pair- 
ing with  one  male,  whilst  others 
are  fighting  for  her,  518;  male, 
attracted  by  the  voice  of  the  fe- 
male, 522;  male,  odor  emitted  by, 
525. 

,    Axis,    sexual   difference   in 

the  color  of  the,  532. 

-,     fallow,     different     colored 


herds   of,    535. 

,  Mantchurian,  541. 

,  Virginian,  541;  color  of  the, 

not   affected  by  castration,   530; 

colors  of,  531. 
Deerhound,  Scotch,  greater  size  of 

the  male,  232,  512. 
Defensive  organs     of     mammals, 

513. 
De  Geer,    C,   on  a  female  spider 

destroying  a  male,  269. 
Dekay,   Dr.,   on   the   bladder-nose 

seal    524. 
Delorenzi,    G.,    division   of    malar 

bone,   37. 
Demerara,   yellow  fever  in,  189. 
Dendrocygna,  459. 
Dendrophila  frontalis,    young   of, 

483.  ,     ., 

Denison,  Sir  W.,  manner  of  rid- 
ding themselves  of  vermin 
among  the  Australians,  55;  ex- 
tinction of  Tasmanians,  179. 

Denny,  H.,  on  the  lice  of  domestic 
animals,   165. 

Dermestes  murinus,  stridulation 
of,  299. 

Descent  traced  through  the 
mother  alone,  583. 

Deserts,  protective  coloring  of 
animals  inhabiting,  485. 

Desmarest,  on  the  absence  of 
suborbital  pits  in  Antilope  sub- 
gutturosa,  525:  on  the  whiskery 
of  Macacus,  527;  on  the  color  of 
the  opossum,  530;  on  the  colors 
of  the  sexes  of  Mus  minutus, 
529;  on  the  coloring  of  the  oce- 
lot,  529;   on   the  colors  of  seals. 


530;  on  Antilope  caama,  531;  on 
the  colors  of  goats,  531;  on  sex- 
ual difference  of  color  in  Ateles 
marginatus,  532;  on  the  mandrill, 
534;  on  Macacus  cynomolgus, 
553. 

Desmoulins,  on  the  number  of 
species  of  man,  170;  on  the 
musk-deer,  525. 

Desor,  on  the  imitation  of  man  by 
monkeys,   71. 

Despine,  P.,  on  criminals  destitute 
of   conscience,    112. 

Development,  embryonic,  of  man, 
9,   11;   correlated,  420. 

Devil,  not  believed  in  bj'  the  Fue- 
gians,  92. 

Devil-crab,  265. 

Devonian,  fossil  insect  from  the, 
286. 

Dewlaps,  of  cattle  and  antelopes, 
527. 

Diadema,  sexual  differences  of 
coloring  in  the  species  of,  306. 

Diamond-beetles,  bright  colors  of. 
291. 

Diastema,  occurrence  of,  in  man, 
34. 

Diastylidse,  proportion  of  the 
sexes  in,  251. 

Dicrurus,  racket-shaped  feathers 
in,   379;   nidification   of,   447. 

macrocercus,  change  of  plu- 
mage in,   454. 

Didelphis  opossum,  sexual  differ- 
ence in  the  color  of,  529. 

Differences,  comparative,  between 
different  species  of  birds  of  the 
same  sex,  464. 

Digits,  supernumerary,  more  fre- 
quent in  men  than  in  women, 
219;  supernumerary,  inheritance 
of,  227;  supernumerary,  early  de- 
velopment of,  232. 

Dimorphism  in  females  of  water- 
beetles,  273;  in  Neurothemis  and 
AgrioH;    288. 

Diodorus,  on  the  absence  of  beard 
in  the  natives   of   Ceylon,   555. 

Dipelicus  Cantori,  sexual  differ- 
ences of,  294. 

Diplopoda,  prehensile  limbs  of  the 
male,  271. 

Dipsas  cynodon,  sexual  difference 
in  the  color  of,  347. 

Diptera,   278. 

Disease,  generated  by  the  contact 
of  distinct  peoples,  178. 

Diseases  common  to  man  and  the 
lower  animals,  7;  difference  of 
liability  to,  in  different  races  ot 
men,  163;  new,  effects  of,  upon 
savages,  178;  sexually  limited, 
232 

Display,  coloration  of  Lepidoptera 
for  31:  of  plumage  by  male 
birds,  390,  396.  ^  ^^ 

Distribution,  wide,  of  man,  46; 
geographical,  as  evidence  of  spe- 
cific distinctness  in  man,  165. 

Disuse,  effects  of,  in  producing  ru- 


INDEX. 


631 


dimentary  organs,  11;  and  use 
of  parts,  effects  of,  31;  of  parts, 
influence  of,  on  the  races  of 
men,    192. 

Divorce,  freedom  of,  among  the 
Charruas,  592. 

Dixon,  E.  S.,  on  the  pairing  of  dif- 
ferent species  of  geese,  409;  on 
the  courtship  of  peafowl,  414. 

Dobrizhoffer,  on  the  marriage- 
customs  of  the  Abipones,  593. 

Dobson,  Dr.,  on  the  Cheiroptera, 
214;  scent-glands  of  bats,  524; 
frugivorous  bats,  529. 

Dogs,  suffering  from  Tertian 
ague,  8;  memory  of,  72;  dream- 
ing, 72;  diverging  when  drawing 
sledges  over  thin  ice,  73;  exer- 
cise of  reasoning  faculties  by, 
75;  domestic,  progress  of  in 
moral  qualities,  78;  distinct 
tones  uttered  by,  82;  parallelism 
between  his  affection  for  his 
master  and  religious  feeling,  93; 
sociability  of  the,  97;  sympathy 
of,  with  a  sick  cat,  100;  sympa- 
thy of,  with  his  master,  100; 
their  possession  of  conscience, 
100;  possible  use  of  the  hair  on 
the  fore-legs  of  the,  147;  races 
of  the,  172;  numerical  proportion 
of  male  and  female  births  in, 
240;  sexual  affection  between  in- 
dividuals of,  519;  howling  at  cer- 
tain notes,  563;  rolling  in  car- 
rion, 525. 

Dolichocephalic  structure,  possi- 
ble cause  of,  54. 

Dolphins,  nakedness  of,  54. 

Domestic  animals,  races  of,  172; 
change  of  breeds  of,  590. 

Domestication,  influence  of,  in  re- 
moving the  sterility  of  hybrids, 
168. 

D'Orbignj'',  A.,  on  the  influence  of 
dampness  and  dryness  on  the 
color  of  the  skin,  188;  on  the 
Yuracaras,   574. 

Dotterel,  477. 

Doubleday,  E.,  on  sexual  differ- 
ences in  the  wings  of  butterflies, 
274. 

-,  H.,  on  the  proportion  of  the 


sexes  in  the  smaller  moths,  247; 
males  of  Lasiocampa  quercus 
and  on  the  attraction  of  the  Sa- 
turnia  carpini  by  the  female, 
252;  on  the  proportion  of  the 
sexes  in  the  Lepidoptera,  248;  on 
the  ticking  of  Anobium  tessel- 
atum,  307;  on  the  structure  of 
Ageronia  feronia,  305;  on  white 
butterflies  alighting  upon  paper, 
314. 

Douglas,  J.  W.,  on  the  sexual  dif- 
ferences of  the  Hemiptera,  278; 
on  the  colors  of  British  Homop- 
tera,  280. 

Down,  of  birds,  385. 

Draco,  gular  appendages  of,  350. 


Dragonet,  Gemmeors,  332. 
Dragon-flies,    caudal     appendages 

of  male,  274;  relative  size  of  the 

sexes   of,    276;    difference   in   the 

sexes  of,  287;  want  of  pugnacity 

by  the  male,  288. 
Drake,   breeding  plumage  of  the, 

387. 
Dreams,   72;   a  possible  source   of 

the  belief  in  spiritual   agencies, 

91. 
Drill,    sexual    difference    of   color 

in  the,  533. 
Dromseus    irroratus,  472. 
Dromoleea,    Saharan    species     of, 

450. 
Drongo  shrike,  454. 
Drongos,   racket-shaped     feathers 

in  the  tails  of,  379,   387. 
Dryness,  of  climate,  supposed  in- 
fluence  of,   on   the  color  of  the 

skin,   188. 
Dryopithecus,  151. 
Duck,    harlequin,    age   of    mature 

plumage  in  the,  478;  breeding  in 

immature  plumage,  479. 
,    long-tailed,    preference     of 

male,  for  certain  females,  415. 
-,  pintail,  pairing  with  a  widg- 


eon, 409. 


,  voice   of  the,   369;      pairing 

with  a  shield-drake,  409;  imma- 
ture plumage  of  the,  461. 

-,   wild,    sexual  differences   in 


the,  214;  speculum  and  male 
characters  of,  231;  pairing  with 
a  pintail  drake,  410. 

Ducks,  wild,  becoming  polygam- 
ous under  partial  domestication, 
215;  dogs  and  cats  recognized 
by,  406. 

Dufosse,  Dr.,  sounds  produced  by 
flsh,  343. 

Dugong,  nakedness  of,  54;  tusks 
of,  498. 

Dujardin,  on  the  relative  size  of 
the  cerebral  ganglia  in  insects, 
52. 

Duncan,  Dr.,  on  the  fertility  of 
early  marriages,  134;  compara- 
tive health  of  married  and  sin- 
gle, 136. 

Dupont,  M.,  on  the  occurrence  of 
the  supra-condyloid  foramen  in 
the  humerus  of  man,  21. 

Durand,  J.  P.,  on  causes  of  varia- 
tion, 29. 

Dureau  de  la  Malle,  on  the  songs 
of  birds,  84;  on  the  acquisition 
of  an  air  by  blackbirds,  366. 

Dutch,  retention  of  their  color  by 
the,  in  South  Africa,  188. 

Duty,  sense  of,  94. 

Duvaucel,  female  Hylobates 
washing  her  young,  68. 

Dyaks,  pride  of,  in  mere  homi- 
cide,  113. 

Dynastes,  large  size  of  males  of, 
276. 

Dynastini,   stridulation  of,   300. 


632 


INDEX. 


Dytisctis,  dimorphism  of  females 
of,  273;  g-rooved  elytra  of  the  fe- 
male,  273. 


E 


Eagle,  young-  Cercopithecus  res- 
cued from,  by  the  troop,  98. 

,    white-headed,    breeding    in 

immature  plumage,  479. 

Eagles,  golden,  new  mates  found 
by,  403. 

Ear,  motion  of  the,  13;  external 
shell  of  the,  usele&s  in  man,  14; 
rudimentary  point  of  the,  in 
man,  14. 

Ears,  more  variable  in  men  than 
women,  219;  piercing  and  orna- 
mentation of  the,  570. 

Earwigs,   parental  feeling  in,  102. 

Echidna,  152. 

Echini,  bright  colors  of  some,  257. 

Echinodermata,  absence  of  sec- 
ondary sexual  characters  in,  257. 

Echis  carinata,  349. 

Ecker,  figure  of  the  human  em- 
bryo, 10;  on  the  development  of 
the  gyri  and  sulci  of  the  brain, 
201;  on  the  sexual  differences  in 
the  pelvis  in  man,  552;  on  the 
presence  of  a  sagittal  crest  in 
Australians,  553. 

Edentata,  former  wide  range  of, 
in  America,  165;  absence  of  sec- 
ondary sexual  characters  in,  214. 

Edolius,  racket-shaped  feathers 
in,  379. 

Edwards,  Mr.,  on  the  proportion 
of  the  sexes  in  North  American 
species  of  Papilio,  246. 

Eels,   hermaphroditism  of,   157. 

Egerton,  Sir  P.,  on  the  use  of  the 
antlers  of  deer,  506;  on  the  pair- 
ing of  red  deer,  518;  on  the  bel- 
lowing of  stags,   521. 

Eggs,  hatched  by  male  fishes,  341. 

Egret,  Indian,  sexes  and  young 
of,  481. 

Egrets,  breeding  plumage  of,  385; 
white,  488. 

Ehrenberg,  on  the  mane  of  the 
male   Hamadryas   baboon,   517. 

Ekstrom,  M.,  on  Harelda  glacial- 
is,   415. 

Elachista  rufocinerea,  habits  of 
male,  248. 

Eland,  development  of  the  horns 
of  the,  229. 

Elands,  sexual  differences  of  color 
in,  531. 

Elaphomyia,  sexual  differences  in, 

277. 
Elaphrus   uliginosus,    stridulation 

of,  300. 
Elaps,  349. 
Elateridse,      proportions      of      the 

sexes  in,  249. 
Elaters,  luminous,  275. 


Elephant,  152;  rate  of  increase  of 
the,  45;  nakedness  of  the,  55;  In- 
dian, forbearance  to  his  keeper, 
101;  polygamous  habits  of  the, 
213;  pugnacity  of  the  male,  497; 
tusks  of,  498,  499,  503,  510;  In- 
dian, mode  of  fighting  of  the, 
509;  male,  odor  emitted  by  the, 
524;  attacking  white  or  gray 
horses,  535. 

Elevation  of  abode,  modifying  in- 
fluence of,  33. 

Elimination  of  inferior  individu- 
als, 133. 

Elk,  503;  winter  change  of  the, 
537. 

,   Irish,  horns  of  the,  511. 

Ellice  Islands,  beards  of  the  na- 
tives, 555,  575. 

Elliot,  R.,  on  the  numerical  pro- 
portion of  the  sexes  in  young 
rats,  241;  on  the  proportion  of 
the  sexes  in  sheep,  241. 

,    D.    G.,    on    Pelecanus    ery- 

throrhynchus,  384. 

-,  Sir  W.,  on  the  polygamous 


habits  of  the  Indian  wild  boar, 
213. 

Ellis,  on  the  prevalence  of  infan- 
ticide in  Polynesia,  586. 

Elphinstone,  Mr.,  on  local  differ- 
ences of  stature  among  the  Hin- 
doos, 30;  on  the  difficulty  of  dis- 
tinguishing the  native  races  of 
India,  163. 

Elytra,  of  the  females  of  Dytis- 
cus,  Acilius,  Hydroporus,  273. 

Emberiza,  characters  of  young, 
459. 

miliaria,  459. 

schoenicius,  407:  head-feath- 
ers of  the  male,  393. 

Embryo  of  man,  9,  10;  of  the  dog, 
10. 

Embryos  of  mammals,  resem- 
blance of  the,  24. 

Emigration,   133. 

Emotions  experienced  by  the 
lower  animals  in  common  with 
man,  67;  manifested  by  animals, 
69. 

Emperor  moth,  312. 

Emulation   of  singing-birds,   364. 

Emu,  sexes  and  incubation  of, 
472. 

Endurance,    estimation  of,  115. 

Energy,  a  characteristic  of  men, 
559. 

England,  numerical  proportion  of 
male  and  female  births  in,  237. 

Engleheart,  Mr.,  on  the  finding 
of  new  mates  by  starlings,  403. 

English,  success  of,  as  colonists, 
138. 

Engravers,    short-sighted,   32. 

Entomostraca,  265. 

EntOzoa,  difference  of  color  be- 
tween the  males  and  females  of 
some,  257. 

Envy,  persistence  of,  109. 


INDEX. 


633 


Eocene  period,  possible  divergence 

of  man  during  the,  151. 
Eolidse,     colors    of,    produced    by 
the  biliary  glands,  258. 

Epeira   nigra,    small    size    of    the 

male  of,  269. 
Ephemera,   272. 
Ephemeridse,  287. 

Ephippiger  vitium,  stridulating 
organs  of,  283,  284. 

Epicalia,  sexual  differences  of  col- 
oring in  the  species  of,  306. 

Equus  hemionus,  winter  change 
of,   537. 

Erateina,  coloration  of,  312. 

Ercolani,  Prof.,  hermaphroditism 
in  eels,  157. 

Erect  attitude  of  man,  49,  50. 

Eristalis,  courting  of,  278. 

Eschricht,  on  the  development  of 
hair  in  man,  18;  on  a  lanuginous 
moustache  in  a  female  foetus, 
19;  on  the  want  of  definition  be- 
tween the  scalp  and  the  fore- 
head in  some  children,  147;  on 
the  arrangement  of  the  hair  in 
the  human  foetus,  157;  on  the 
hairiness  of  the  face  in  the  hu- 
man foetus  of  both  sexes,  597, 
598. 

Esmeralda,  difference  of  color  in 
the  sexes  of,  292. 

Esox  lucius,  244. 

reticulatus,  337. 

Esquimaux,  61,  129;  their  belief  in 
the  inheritance  of  dexterity  in 
seal-catching,  32;  mode  of  life 
of,  192. 

Estrelda  amandava,  pugnacity  of 
the  male,   362. 

Eubagis,  sexual  differences  of 
coloring  in  the  species  of,  306. 

Euchirus  longimanus,  sound  pro- 
duced by,   301. 

Eudromias  morinellus,  471. 

Eulampis  jugularis,  colors  of  the 
female,  447. 

Euler,  on  the  rate  of  increase  in 
the  United  States,  43. 

Eumomota  superciliaris,  racket- 
shaped  feathers  in  the  tail  of, 
379. 

Eupetornena  macroura,  colors  of 
the  female,  447. 

Euphema  splendida,  451. 

Euplocamus  erythrophthalmus, 
possession  of  spurs  by  the  fe- 
male, 360. 

Europe,  ancient  inhabitants  of, 
177. 

Europeans,  difference  of,  from 
Hindoos,  187;  hairiness  of,  prob- 
ably due  to  reversion,  596. 

Eurostopodus,  sexes  of,  473. 

Eurygnathus,  different  propor- 
tions of  the  head  in  the  sexes 
of,  274. 

Eustephanus,  sexual  differences 
of  species  of,   356;  young  of,  483. 

Exaggeration  of  natural  charac- 
ters by  man,  577. 


Exogamy,  583,  586. 

Expression,  resemblances  in,  be- 
tween man  and  the  apes,  146. 

Extinction  of  races,  causes  of,  177. 

Eye,  destruction  of  the,  31; 
change  of  position  in,  53;  ob- 
liquity of,  regarded  as  a  beauty 
by  the  Chinese  and  Japanese, 
572. 

Eyebrows,  elevation  of,  13;  devel- 
opment of  long  hairs  in,  18;  in 
monkeys,  147;  eradicated  in 
parts  of  South  America  and  Af- 
rica, 569;  eradication  of,  by  the 
Indians  of  Paraguay,  575. 

Eyelashes,  eradication  of,  by  the 
Indians  of  Paraguay,  575. 

Eyelids,  colored  black,  in  part  of 
Africa,  569. 

Eyes,  pillared,  of  the  male  of 
Chloeon.  272;  difference  in  the 
color  of,  in  the  sexes  of  birds, 
419. 

Eyton,  T.  C,  observations  on  the 
development  of  the  horns  in  the 
fallow-deer,  229. 

Eyzies,  Les,  human  remains  from, 
177. 


F 


Fabre,  M.,  on  the  habits  of  Cer- 
ceris,  289. 

i^'acial  bones,  causes  of  modifica- 
tion of  the,  53. 

Faculties,  diversity  of,  in  the 
same  race  of  men,  25;  inherit- 
ance of,  27;  diversity  of,  in  ani- 
mals of  the  same  species,  27; 
mental,  variation  of,  in  the  same 
species,  64;  of  birds,  405. 

Fakirs,  Indian,  tortures  under- 
gone by,  115. 

Falco  leucocephalus,  479. 

peregrinus,  402,  454. 

tinnunculus,  402. 

Falcon,  peregrine,  new  mate 
found  by,  402. 

Falconer,  H.,  on  the  mode  of 
fighting  of  the  Indian  elephant, 
509;  on  canines  in  a  female  deer, 
510;  on  Hyomoschus  aquaticus, 
542. 

Falkland  Islands,  horses  of,  176. 

Fallow-deer,  different  colored 
herds  of,  535. 

Famines,  frequency  of,  among 
savages,  44. 

Farr,  Dr.,  on  the  structure  of  the 
uterus,  37;  on  the  effects  of  prof- 
ligacy, 134;  on  the  infiuence  of 
marriage  on   mortality,   135,   136. 

Farrar,  F.  W.,  on  the  origin  of 
language,  84;  on  the  crossing  or 
blending  of  languages,  88;  on  the 
absence  of  the  idea  of  God  in 
certain  races  of  men,  91;  on 
early  marriages  of  the  poor,  134; 
on  the  middle  ages,  138. 


634 


INDEX. 


Fashions,  long  prevalence  of, 
among  savages,  571,  578. 

Faye,  Prof.,  on  the  numerical 
proportion  of  male  and  female 
births  in  Norway  and  Russia, 
238;  on  the  greater  mortality  of 
male  children  at  and  before 
birth,  238. 

Feathers,  modified,  producing 
sounds,  372  et  seq.,  444;  elon- 
gated, in  male  birds,  379,  397; 
racket-shaped,  380;  barbless  and 
with  filamentous  barbs  in  cer- 
tain birds,  380;  shedding  of  mar- 
gins of,  388. 

Feeding,  high,  probable  influence 
of,  in  the  pairing  of  birds  of 
different  species,  410. 

Feet,  thickening  of  the  skin  on 
the  soles  of  the,  32;  modification 
of,  in  man,  50. 

Felis  canadensis,  throat-ruff  of, 
516. 

pardalis  and  F.  mitis,  sex- 
ual differences  in  the  coloring 
of,  529. 

Female,  behavior  of  the,  during 
courtship,  217. 

birds,  differences  of,  464. 

Females,  presence  of  rudimentary 

male  organs  in,  159;  preference 
of,  for  certain  males,  210;  pur- 
suit of,  by  males,  216;  occur- 
rence of  secondary  sexual  char- 
acters in,  220;  development  of 
male  characters  by,  222. 

and    males,      comparative 

numbers  of,  209,  210;  comparative 
mortality  of,  while  young,  211. 

Femur  and  tibia,  proportions  of, 
in  the  Aymara  Indians,  33. 

Fenton,  Mr.,  decrease  of  Maories, 
180;  infanticide  amongst  the 
Maories,  253. 

Ferguson,  Mr.,  on  the  courtship 
of  fowls,  412. 

Fertility  lessened  under  changed 
conditions,  183. 

Fertilization,  phenomena  of,  in 
plants,  217;  in  the  lower  animals, 
218. 

Fevers,  immunity  of  Negroes  and 
Mulattoes  from,   188. 

Fiber  zibethicus,  protective  color- 
ing of  it,  537. 

Fick,  H.,  effect  of  conscription  for 
military  service,  131. 

Fidelity  of  savages  to  one  an- 
other, 114;  importance  of,  120. 

Field-slaves,  difference  of,  from 
house-slaves,  191. 

Fijians,  burying  their  old  and 
sick  parents  alive,  99;  estimation 
of  the  beard  among  the,  575;  ad- 
miration of,  for  a  broad  occi- 
put, 577. 

Fiji  Archipelago,  population  of 
the,  169. 

Islands,  beards  of  the  na- 
tives, 555,  575;  marriage-customs 
of  the.  592. 


Filial  affection,   partly  the  result 

of  natural  selection,  102. 
Filum  terminale,  22. 
Finch,   racket-shaped   feathers  in 

the  tail  of  a,   380. 
Finches,    spring   change   of   color 
in,  388;   British,  females  of  the, 
453. 

Fingers,  partially  coherent,  in 
species  of  Hylobates,  49. 

Finlayson,  on  the  Cochin  Chinese, 
573. 

Fire',   use  of,   47,   141,  175. 

Fischer,  on  the  pugnacity  of  the 
male  of  Lethrus  cephalotes, 
297. 

Fish,  eagerness  of  male,  216;  pro- 
portion of  the  sexes  in,  244; 
sounds   produced   by,    343. 

Fishes,  kidneys  of,  represented  by 
Corpora  Wolffiana  in  the  human 
embryo,  11;  male,  hatching  ova 
in  their  mouths,  158;  receptacles 
for  ova  possessed  by,  204;  rela- 
tive size  of  the  sexes  in,  331; 
fresh-water,  of  the  tropics,  339; 
protective  resemblances  in,  340; 
change  of  color  in,  340;  nest- 
building,  341;  spawning  of,  341; 
sounds  produced  by,  343,  561; 
continued  growth  of,  480. 

Flamingo,  age  of  mature  plu- 
mage, 478. 

Flexor  pollicis  longus,  similar  va- 
riation of,  in  man,  41. 

Flint  tools.  141. 

Flints,  difficulty  of  chipping  into 
form,  48. 

Floresuga  mellivora,  437. 

Florida,  Quiscalus  major  in,  224. 

Flounder,  coloration  of  the,  340. 

Flower,  W.  H.,  on  the  abductor 
of  the  fifth  metatarsal  in  apes, 
40;  on  the  position  of  the  Seals, 
146;  on  the  Pithecia  monachus, 
197;  on  the  throat-pouch  of  the 
male  bustard,   368. 

Fly-catchers,  colors  and  nidifica- 
tion  of,  449. 

Foetus,  human,  woolly  covering 
of  the,  19;  arrangement  of  the 
hair   on,   148. 

Food,  influence  of,  upon  stature, 
30. 

Foot,  prehensile  power  of  the,  re- 
tained in  some  savages,  50;  pre- 
hensile, in  the  early  progenitors 
of  man,  156. 

Foramen,  supra-condyloid,  excep- 
tional occurrence  of  in  the  hum- 
erus of  man,  21,  42;  in  the  early 
progenitors  of  man,  156. 

Forbes,  D.,  on  the  Aymara  In- 
dians, 33;  on  local  variation  of 
color  in  the  Quichuas,  191;  on 
the  hairlessness  of  the  Aymaras 
and  Quichuas,  556;  on  the  long 
hair  of  the  Aymaras  and  Qui- 
chuas.  554,   574. 

Forel,  F.,  on  white  young  swans, 
477. 


INDEX. 


635 


Formica  rufa,  size  of  the  cerebral 
ganglia  in,   52. 

Fossils,  absence  of,  connecting 
man  with  the  apes,  152. 

Fowl,  occurrence  of  spurs  in  the 
female,  223;  game,  early,  pug- 
nacity of,  233;  Polish,  early  de- 
velopment of  cranial  peculiari- 
ties of,  234;  variations  in  plum- 
age of,  3S0;  examples  of  corre- 
lated development  in  the,  420; 
domestic,  breeds  and  subbreeds 
of,  454. 

Fowls,  spangled  Hamburgh,  224, 
233;  inheritance  of  changes  of 
plumage  by,  224;  sexual  peculi- 
arities in, transmitted  only  to  the 
same  sex,  225;  loss  of  second- 
ary sexual  characters  by  male, 
226;  Polish,  origin  of  the  crest 
in,  226;  period  of  inheritance  of 
characters  by,  233;  cuckoo,  233; 
development  of  the  comb  in,  233; 
numerical  proportion  of  the 
sexes  in,  242;  courtship  of,  411; 
mongrel,  between  a  black  Span- 
ish cock  and  different  hens, 
421;  penciled  Hamburgh,  differ- 
ence of  the  sexes  in,  441;  Span- 
ish, sexual  differences  of  the 
comb  in,  441;  spurred,  in  both 
sexes,  443. 

Fox,  W.  D.,  on  some  half -tamed 
wild  ducks  becoming  polygam- 
ous, and  on  polygamy  in  the 
guinea-fowl  and  canary-bird, 
215;  on  the  proportion  of  the 
sexes  in  cattle,  241;  on  the  pug- 
nacity of  the  peacock,  360;  on  a 
nuptial  assembly  of  magpies, 
401;  on  the  finding  of  new  mates 
by  crows,  402;  on  partridges  liv- 
ing in  triplets,  404;  on  the  pair- 
ing of  a  goose  with  a  Chinese 
gander,  409. 

Foxes,  wariness  of  young,  in 
hunting  districts,  77;   black,  535. 

Fraser,  C,  on  the  different  colors 
of  the  sexes  in  a  species  of 
Squilla,   267. 

,   G.,   colors  of  Thecla,   309. 

Frere,  Hookham,  quoting  Theog- 
nis  on  selection  in  mankind,  28. 

Fringilla  cannabina,   388. 

ciris,  age  of  mature  plumage 

in,  478. 

cyanea,  age  of  mature  plu- 


mage in,  478. 

leucophrys,  young  of,  481. 

spinus,  410. 

tristis,   change  of   color  in, 


in  spring,  388;  young  of,  480. 

Fringillidse,  resemblance  of  the 
females  of  distinct  species  of, 
464. 

Frogs,  345;  male,  temporary  re- 
ceptacles for  ova  possessed  by, 
204;  ready  to  breed  before  the  fe- 
males, 208;  fighting  of,  345;  vocal 
organs  of,  346. 


Frontal   bone,    persistence   of   the 
suture  in,  38. 

Fruits,  poisonous,  avoided  by  ani- 
mals, 65. 

Fuegians,    130,    140;    difference    of 
stature  among  the,  30;  power  of 
sight  in  the,  32;  skill  of,  in  stone- 
throwing,   48;   resistance  of  the 
to  their  severe  climate,   61,   177 
mental      capacity     of     the,     63 
quasi-religious     sentiments      of 
the,  92;  resemblance  of,  in  men- 
tal  characters,     to     Europeans. 
174;    mode    of    hfe    of    the,    192; 
aversion  of,  to  hair  on  the  face 
575;    said    to    admire    Europear 
women,  576. 

Fulgoridse,  songs  of  the,  279. 

Fur,   whiteness   of,   in  arctic  ani- 
mals, in  winter,  224. 

Fur-bearing       animals,     acquired 
sagacity  of,  77. 


G 


Gallicrex,  sexual  difference  in  the 
color  of  the  irides  in,  419. 

cristatus,  pugnacity  of  male, 

357;  red  caruncle  occurring  in 
the  male  during  the  breeding- 
season,  384. 

Gallinaceae,  frequency  of  polyg- 
amous habits  and  of  sexual  dif- 
ferences in  the,  215;  love-ges- 
tures of,  375;  decomposed  feath- 
ers in,  380;  stripes  of  young, 
459;  comparative  sexual  differ- 
ences between  the  species  of, 
464,   465;  plumage  of,   466. 

Gallinaceous  birds,  weapons  of 
the  male,  358;  racket-shaped 
feathers  on  the  heads  of,   379. 

Gallinula  chloropus,  pugnacity  of 
the  male,  356. 

Galloperdix,  spurs  of,  360;  devel- 
opment of  spurs  in  the  female, 
444. 

Gallophasis,  young  of,  463. 

Gallus  bankiva,  441;  neck-hackles 
of,    387. 

Stanleyi,    pugnacity   of   the 

male,  359. 

Galls,  58. 

Galton,  Mr.,  on  hereditary  gen- 
ius, 27;  gregariousness  and  in- 
dependence in  animals,  101;  on 
the  struggle  between  the  social 
and  personal  impulses,  121;  on 
the  effects  of  natural  selection 
on  civilized  nations,  130;  on  the 
sterility  of  sole  daughters,  132; 
on  the  degree  of  fertility  of  peo- 
ple of  genius,  133;  on  the  early 
marriages  of  the  poor,  134;  on 
the  ancient  Greeks,  137;  on  the 
Middle  Ages,  137;  on  the  prog- 
ress of  the  United  States,  138: 
on  South  African  notions  of 
beauty,  574. 


636 


INDEX. 


Gammarus,  use  of  the  chelae  of, 
265. 

marinus,  266. 

Gannets,  white  only  when  ma- 
ture, 488. 

Ganoid  fishes,  154,  160. 

Gaour,  horns  of  the,  501. 

Gap  between  man  and  the  apes, 
152. 

Gaper,  sexes  and  young  of,  481. 

Gardner,  on  an  example  of  ra- 
tionality in  a  Gelasimus,  267. 

Garrulus  glandarius,  402. 

Gartner,  on  sterility  of  hybrid 
plants,  168. 

Gasteropoda,  259;  pulmoniferous, 
courtship  of,  259.  ^ 

Gasterosteus,  216;  nidification  of, 
341 

. 1  leiurus,  328,  337,  341. 

trachurus,  328. 

Gastrophora,  wings  of,  brightly 
colored    beneath,    312. 

Gauchos,  want  of  humanity 
among  the,  119. 

Gaudry,  M.,  on  a  fossil  monkey, 
150. 

Gavia,  seasonal  change  of  plu- 
mage in,  488. 

Geese,  clanging  noise  made  by, 
364;  pairing  of  different  species 
of,  409;  Canada,  selection  of 
mates  by,   411. 

Gegenbaur,  C,  on  the  number  of 
digits  in  the  Ichthyopterygia, 
35-  on  the  hermaphroditism  of 
the  remote  progenitors  of  the 
vertebrata,  157;  two  types  of 
nipple  in  mammals,  158. 

Gelasimus,  proportions  of  the 
sexes  in  a  species  of,  251;  use  ot 
the  enlarged  chelse  of  the  male, 
264;  pugnacity  of  males  of,  265; 
rational  actions  of  a,  267;  dif- 
ference of  color  in  the  sexes  of 
a  species  of,  268. 

G^mmules,    dormant   in    one    sex, 

226. 
Genius,  27;   hereditary,  559. 

,  fertility  of  men  and  women 

of,   133. 

Geoff  roy-Saint-Hilaire,  Isid.,  on 
the  recognition  of  women  by 
ntiale  quadrumana,  8;  on  mon- 
strosities, 29;  coincidences  of  ar- 
rested development  with  poly- 
dactylism,  35;  on  animal-like 
anomalies  in  the  human  struc- 
ture, 38;  on  the  correlation  of 
monstrosities,  42;  on  the  distrib- 
ution of  hair  in  man  and  mon- 
keys, 55;  on  the  caudal  verte- 
brae of  monkeys,  56;  on  corre- 
lated variability,  58;  on  the 
classification  of  man,  143;  on  the 
long  hair  on  the  heads  of  species 
of  Semnopithecus,  146;  on  the 
hair  in  monkeys,  148;  on  the  de- 
velopment of  horns  in  female 
deer,  500;  and  F.  Cuvier,  on  the 


mandrill,  534;  on  Hylobates,  553, 
554. 
Geographical  distribution,  as  evi- 
dence of  specific  distinctions  in 
man,  165. 
Geometrae,    brightly     colored     be- 
neath, 312. 
Geophagus,    frontal   protuberance 
of   male,   366,    341;    eggs   hatched 
by   the   male,    in   the   mouth   or 
branchial  cavity,  341. 
Georgia,  change  of  color  in  Ger- 
mans settled  in,  191. 
Geotrupes,  stridulation  of,  300,  301. 
Gerbe,  M.,  on  the  nest-building  of 
Crenilabrus  massa  and  C.   mel- 
ops,  341. 
Gerland,  Dr.,  on  the  prevalence  of 
infanticide,   113,    372,   587;   on  the 
extinction  of  races,  177. 
Gervais,  P.,   on  the  hairiness     of 
the  gorilla,  55;  on  the  mandrill, 
534. 
Gesture-language,    174. 
Ghost-moth,  sexual  difference    of 

color  in  the,  313. 
Gibbs,    Sir   D.,    on   differences    of 
the   voice   in   different   races  of 
men,  561. 
Gibbon,  Hoolock,  nose  of,  146. 
Gibbons,  voice  of,  522. 
Gill,   Dr.,   male  seals  larger  than 
females,    214;    sexual   differences 
in  seals,  511. 
Giraffe,    its    mode    of   using     the 
horns,   504;   mute,   except  in   the 
rutting  season,  521. 
Girard,    M.,    disputes     descent   of 
vertebrates  from  Ascidians,  155; 
color  of  sponges  and  Ascidia^ns, 
258;  musky  odor  of  Sphinx,  305. 
Giraud-Teulon,    on   the  cause     of 

short  sight,   32. 
Glanders,     communicable  to  man 

from  the  lower  animals,  6. 
Glands,  odoriferous,  in  mammals, 

524,  525. 
Glareola,    double   moult   in,    385. 
Glomeris   limbata,     difference     of 

color  in  the  sexes  of,  270. 
Glow-worm,      female,      apterous, 

204;  luminosity  of  the,  275. 
Gnats,     dances    of,    278;    auditory 

powers  of,  563. 
Gnu,     sexual     differences    in   the 

color  of  the,  531. 
Goat,   male,   wild,   falling  on     his 
horns,    503;    male,    odor    emitted 
by,  524;  male,  wild,  crest  of  the, 
526;  Berbura,  mane,  dewlap,  &c., 
of    the   male,    528;    Kemas,    sex- 
ual difference  in  the  color  of  the, 
531. 
Goats,    sexual    differences    in   the 
horns  of,  225;  horns  of,  229,   501; 
mode  of  fighting  of,  504;  domes- 
tic,  sexual      differences  of,   late 
developed,   232;   beards   of,    526. 
Goatsucker,  Virginian,  pairing  of 
the,  362. 


INDEX. 


637 


CrObies,  nidification  of,  341. 

God,  want  of  the  idea  of,  in  some 
races  of  men,   91. 

Godron,  M.,  on  variability,  29;  on 
difference  of  stature,  30;  on  the 
want  of  connection  between  cli- 
mate and  the  color  of  the  skin, 
188;  on  the  odor  of  the  skin,  193; 
on   the   color   of  infants,   553. 

Goldfinch,  366,  388;  proportion  of 
the  sexes  in  the  243;  sexual  dif- 
ferences of  the  beak  in  the,  356; 
courtship  of   the,   396. 

,  North  American,  young  of, 

480. 

Gold-fish,  339. 

Gomphus,  proportions  of  the 
sexes  in,  250;  difference  in  the 
sexes  of,  287. 

Gonepteryx  Rhamni,  309;  sexual 
difference  of  color  in,  319. 

Goodsir,  Prof.,  on  the  affinity  of 
the  lancelet  to  the  ascidians, 
155. 

Goosander,  young-  of,  462. 

Goose,  Antarctic,  colors  of  the, 
488. 

,  Canada,  pairing  with  a  Ber- 

nicle  gander,  409. 

-,  Chinese,  knob  on  the  beak 


of  the,  420. 

,  Egyptian,  360. 

-,    Sebastopol,      plumage      of, 


380. 


-,  Snow-,  whiteness  of  the, 
488. 

,  Spur-winged,  360. 

Gorilla,  556;  semi-erect  attitude  of 
the,  50;  mastoid  processes  of  the, 
51;  direction  of  the  hair  on  the 
arms  of  the,  147;  manner  of  sit- 
ting, 147;  supposed  to  be  a  kind 
of  mandrill,  173;  polygamy  of 
the,  212,  585;  voice  of  the,  522; 
cranium  of,  553j  fighting  of 
Tnfilp     557 

Gosse,'P.  H.,  on  the  pugnacity  of 
the  male  Humming-bird,   356. 

,    M.,    on   the   inheritance   of 

artificial  modifications  of  the 
skull,  598. 

Gould,  B.  A.,  on  variation  in  the 
length  of  the  legs  in  man,  25; 
measurements  of  American  sol- 
diers, 30,  31;  on  the  proportions 
of  the  body  and  capacity  of  the 
lungs  in  different  races  of  men, 
163;  on  the  inferior  vitality  of 
mulattoes,  167. 

,  J.,  on  migration  of  swifts, 

104;  on  the  arrival  of  male 
snipes  before  the  females,  208; 
on  the  numerical  proportion  of 
the  sexes  in  birds,  242;  on  Neo- 
morpha  Grypus,  356;  on  the  spe- 
cies of  Eustephanus,  356;  on  the 
Australian  musk-duck,  355;  on 
the  relative  size  of  the  sexes  in 
Briziura  lobata  and  Cincloram- 
phus  cruralis,  358;  on  Lobivanel- 


lus  lobatus,  362;  on  the  habits  of 
Menura  Alberti,  366;  on  the  rar- 
ity of  song  in  brilliant  birds, 
366;  on  Selasphorus  platycercus, 
373;  on  the  Bower-birds,  376,  401; 
on  the  ornamental  plumage  of 
the  Humming-birds,  382;  on  the 
moulting  of  the  ptarmigan,  386; 
on  the  display  of  plumage  by 
the  male  Humming-birds,  389; 
on  the  shyness  of  adorned  male 
birds,  397;  on  the  decoration  of 
the  bowers  of  Bower-birds,  408; 
on  the  decoration  of  their  nests 
by  Humming-birds,  408;  on  vari- 
ation in  the  genus  Cynanthus, 
417;  on  the  color  of  the  thighs  in 
a  male  parakeet,  418;  on  Ur- 
osticte  Benjamini,  436,  437;  on 
the  nidification  of  the  Orioles, 
448;  on  obscurely-colored  birds 
building  concealed  nests,  448;  on 
trogons  and  kingfishers,  451;  on 
Australian  parrots,  451;  on  Aus- 
tralian pigeons,  452;  on  the 
moulting  of  the  ptarmigan,  456; 
on  the  immature  plumage  of 
birds,  460  et  seq.;  on  the  Aus- 
tralian species  of  Turnix,  469;  on 
the  young  of  Aithurus  polytmus, 
483;  on  the  colors  of  the  bills  of 
toucans,  487;  on  the  relative  size 
of  the  sexes  in  the  marsupials 
of  Australia,  511;  on  the  colors 
of  the  Marsupials,  529. 

Goureaux,  on  the  stridulation  of 
Mutilla  europaea,  291. 

Gout,   sexually  transmitted,  232. 

Graba,  on  the  Pied  Ravens  of  the 
Feroe  Islands.  418;  variety  of 
the  Guillemot,  418. 

Gradation  of  secondary  sexual 
characters  in  birds,  424. 

Grallatores,  absence  of  secondary 
sexual  characters  in,  215;  double 
moult  in  some,  385. 

Grallina,  nidification  of,  448. 

Grasshoppers,  stridulation  of  the, 
283. 

Gratiolet,  Prof.,  on  the  anthropo- 
morphous apes,  149;  on  the  evo- 
lution of  the  anthropomorphous 
apes,  173;  on  the  difference  in 
the  development  of  the  brains 
of  apes  and  of  man,  199. 

Gray,  Asa,  on  the  gradation  of 
species  among  the  Compositee, 
171. 

,  J.  E.,  on  the  caudal  verte- 
brae of  monkeys,  56;  on  the 
presence  of  rudiments  of  horns 
in  the  female  of  Cervulus  mos- 
chatus,  500;  on  the  horns  of 
goats  and  sheep,  501;  on  the 
beard  of  the  ibex,  526;  on  the 
Berbura  goat,  528;  on  sexual  dif- 
ferences in  the  coloration  of 
Rodents,  529;  ornaments  of  male 
sloth,   529;   on  the  colors  of  the 


638 


INDEX. 


Elande,  531:  on  the  Sing-sine: 
antelope,  531;  on  the  colors  of 
goats,  531;  on  the  hog-deer,  541. 
"Greatest  happiness  principle," 
116,   117. 

Greeks,  ancient,  137. 

Green,  A.  H.,  on  beavers  fighting, 
496;  on  the  voice  of  the  beaver, 
523. 

Greenfinch,  selected  by  a  female 
canary,  410. 

Greg,  W.  R.,  on  the  effects  o^  nat- 
ural selection  on  civilized  na- 
tions, 130;  on  the  early  mar- 
riages of  the  poor,  134;  on  the 
Ancient  Greeks,    137. 

Grenadiers,   Prussian,   28. 

Grey,  Sir  G.,  on  female  infanti- 
cide in  Australia,  587. . 

Greyhounds,  numerical  propor- 
tion of  the  sexes  in,  211,  212; 
numerical  proportion  of  male 
and  female  births  in,  240,   254. 

Grouse,  red,  monogamous,  215; 
nugnacity  of  young  male,  362; 
producing  a  sound  by  scraping 
their  wings  upon  the  ground, 
371;  duration  of  courtship  of, 
400;  colors  and  nidification  of. 
499. 

Gruber,  Dr.,  on  the  occurrence  of 
the  supra-condyloid  foramen  in 
the  humerus  of  man,  21;  on  di- 
vision of  malar  bone,  37;  stridu- 
lation  of  locust,  282. 

Grus  americanus,  age  of  mature 
plumage  in,  478;  breeding  in  im- 
mature plumage,  479. 

Virgo,  trachea  of,  370. 

Gryllus  campestris,  281;  pugnacity 
of  male,  286. 

domesticus,  282. 

Grypus,  sexual  differences  in  the 
beak  in^  356. 

Guanacoes,  battles  of,  496;  canine 
teeth  of,  510. 

Guanas,  strife  for  women  among 
the,  557;  polyandry  among  the, 
588. 

Guanche  skeletons,  occurrence  of 
the  supra-condyloid  foramen  in 
the  humerus  of,  22. 

Guaranys,  proportion  of  men  and 
women  among,  238;  color  of  new- 
born children  of  the,  552;  beards 
of  the,  556. 

Guenee,  A.,  on  the  sexes  of  Hy- 
perythra,  246. 

Guilding,  L.,  on  the  stridulation 
of  the  Locustidae,  280. 

Guillemot,  variety  of  the,  418. 

Guinea,  sheep  of,  with  males  only 
horned,  230. 

Guinea-fowl,  monogamous,  215; 
occasional  polygamy  of  the,  215; 
markings  of  the,  423. 

Guinea-pigs,  inheritance  of  the  ef- 
fects of  operations  by,  597. 

Gulls,  seasonal  change  of  plumage 
in,  488;  white,  488. 


Gunther,  Dr.,  on  paddle  of  Cera- 
todus,  35;  on  hermaphroditism  in 
Serranus,  157;  on  male  fishes 
hatching  ova  in  their  mouths, 
158,  341;  on  mistaking  infertile 
female  fishes  for  males,  244;  on 
the  prehensile  organs  of  male 
Plagiostomous  fishes,  327; 
spines  and  brushes  on  fishes, 
328;  on  the  pugnacity  of  the 
male  salmon  and  trout,  328;  on 
the  relative  size  of  the  sexes  in 
fishes,  331;  on  sexual  differences 
in  fishes,  334  et  seq. ;  on  the  gen- 
us Callionymus,  334;  on  a  pro- 
tective resemblance  in  a  pipe- 
fish, 340;  on  the  genus  Solencs- 
toma,  342;  on  the  coloration  of 
frogs  and  toads,  345;  combats  of 
Testudo  elegans,  347;  on  the  sex- 
ual differences  in  the  Ophidia, 
347;  on  differences  of  the  sexes 
of  lizards,  350  et  seq. 

Gynanisa  Isis,  ocellated  spots  of, 
422. 

Gypsies,  uniformity  of,  in  vari- 
ous parts  of  the  world,  188. 


H 


Habits,  bad,  facilitated  by  famil- 
iarity, 119;  variability  of  the 
force  of,  125. 

Hackel,  E.,  on  the  origin  of  man, 
2;  on  rudimentary  characters, 
11;  on  death  caused  by  inflam- 
mation of  the  vermiform  ap- 
pendage, 20;  on  the  canine  teeth 
in  man,  39;  on  the  steps  by 
which  man  became  a  biped,  50; 
on  man  as  a  member  of  the  Cat- 
arrhine  group,  151;  on  the  posi- 
tion of  the  Lemuridae,  153;  on 
the  genealogy  of  the  Mam- 
malia, 153;  on  the  lancelet,  154; 
on  the  transparency  of  pelagic 
animals,  258;  on  the  musical 
pov/ers  of  women,  567. 

Hagen,  H.,  and  Walsh,  B.  D.,  on 
American  Neuroptera,  251. 

Hair,  development  of,  in  man,  18; 
character  of,  supposed  to  be  de- 
termined by  light  and  heat,  31; 
distribution  of,  in  man,  55,  594; 
possibly  removed  for  ornament- 
al purposes,  55;  arrangement 
and  direction  of,  146,  147;  of  the 
early  progenitors  of  man,  155; 
different  texture  of,  in  distinct 
races,  163;  and  skin,  correlation 
of  color  of,  192:  development  of, 
in  mammals,  526;  management 
of,  among  different  peoples,  569: 
great  length  of,  in  some  North 
American  tribes,  574;  elongation 
of  the,   on  the  human  head,  598. 

Hairiness,  difference  of,  in  the 
sexes  in  man,  554;  variation  of, 
in  races  of  men.  554. 


INDEX. 


639 


Hairs  and  excretory  pores,  nu- 
merical relation  of,  in  sheep,  193. 

Hairy  family,  Siamese,  596. 

Halbertsma,  Prof.,  hermaphrodit- 
ism in  Serranus,  157. 

Hamadryas  baboon,  turning  over 
stones,  98;  mane  of  the  male, 
517. 

Hamilton,  C,  on  the  cruelty  of 
the  Kaffirs  to  animals,  114;  on 
the  engrossment  of  the  women 
by  the  Kaffir  chiefs,  590. 

Hammering,  difficulty  of,  47. 

Hancock,  A.,  on  the  colors  of  the 
nudibranch    Mollusca,    258,   261. 

Hands,  larger  at  birth,  in  the 
children  of  laborers,  32;  struc- 
ture of,  in  the  quadrumana,  48; 
and  arms,  freedom  of,  indirectly 
correlated  with  diminution  of 
canines,  51. 

Handwriting,   inherited,   86. 

Handyside,     Dr.,     supernumerary 

mammae  in  men,  35. 

Harcourt,  E.  Vernon,  on  Fringilla 
cannabina,  388. 

Harelda  glacialis,   415. 

Hare,  protective  coloring  of  the, 
537. 

Hares,  battles  of  male,  496. 

Harlan,  Dr.,  on  the  difference  be- 
tween field-  and  hovise-slaves, 
191. 

Harris,  J.  M.,  on  the  relation  of 
complexion  to  climate,  190. 

,  T.  W.,  on  the  Katy-did  lo- 
cust, 280;  on  the  stridulation  of 
the  grasshoppers,  284;  on  CEcan- 
thus  nivalis,  287;  on  the  coloring 
of  Lepidoptera,  311;  on  the  col- 
oring of  Saturnia  lo,  312. 

Harting,  spur  of  the  Ornithorhyn- 
chus,   498. 

Hartman,  Dr.,  on  the  singing  of 
Cicada  septemdecim,   279. 

Hatred,  persistence  of,  109. 

Haughton,  S.,  on  a  variation  of 
the  flexor  poUicis  longus  in  man, 
41. 

Hawks,  feeding  orphan  nestling, 
404. 

Hayes,  Dr.,  on  the  diverging  of 
sledge-dogs  on  thin  ice,  73. 

Haymond,  R.,  on  the  drumming 
of  the  male  Tetrao  umbellus, 
371;  on  the  drumming  of  birds, 
371. 

Head,  altered  position  of,  to  suit 
the  erect  attitude  of  man,  53; 
hairiness  of,  in  man,  55;  pro- 
cesses of,  in  male  beetles,  292; 
artificial  alterations  of  the  form 
of  the,  577. 

Hearne,  on  strife  for  women 
among  the  North  American  In- 
dians, 556;  on  the  North  Ameri- 
can Indians'  notion  of  female 
beauty,  572;  repeated  elope- 
ments of  a  North  Amei'ican 
woman,  592. 

Heart,  in  the  human  embryo,  9. 


Heat,  supposed  effects  of,  31. 

Hectocotyle,  260. 

Hedge  warbler,  467;  young  of  the, 
475. 

Heel,  small  projection  of,  in  the 
Aymara  Indians,  33. 

Hegt,  M.,  on  the  development  of 
the    spurs     in    peacocks,  230. 

Heliconidse,  305;  mimicry  of,  by 
other  butterflies,  319. 

Heliopathes,  stridulation  peculiar 
to  the  male,  302. 

Heliothrix  auriculata,  young  of, 
462. 

Helix  pomatia,  example  of  in- 
dividual attachment  in,  259. 

Hellins,  J.,  proportions  of  sexes 
of  Lepidoptera  reared  by,  249. 

Helmholtz,  on  pleasure  derived 
from  harmonies,  90;  on  the  vi- 
bration of  the  auditory  hairs  of 
Crustacea,  563;  the  physiology  of 
harmony,  563. 

Hemiptera,  278. 

Hemitragus,  beardless  in  both 
sexes,   526. 

Hemsbach,  M.  von,  on  medial 
mamma  in  man,  35. 

Hepburn,  Mr.,  on  the  autumn 
song  of  the  water-ouzel,  365. 

Hepialus  humuli,  sexual  differ- 
ence of  color  in  the,  313. 

Herbs,  poisonous,  avoided  by  ani- 
mals, 64. 

Hermaphroditism  of  embryos,  157. 

Herodias  bubulcus,  vernal  moult 
of,  387. 

Heron,  Sir  R.,  on  the  habits  of 
peafowl,  413,  414,  437. 

love-gestures  of  a,  375. 

Herons,  decomposed  feathers  in, 
380;  breeding  plumage  of,  386; 
young  of  the,  475;  some- 
times dimorphic,  479;  continued 
growth  of  crest  and  plumes  in 
the  males  of  some,  480;  change 
of  color  in  some,  489. 

Hesperomys  cognatus,   562. 

Hetaerina,  proportion  of  the  sexes 
in,  250;  difference  in  the  sexes 
of,  287. 

Heterbcerus,    stridulation   of,   299. 

Hewitt,  Mr.,  on  a  game-cock  kill- 
ing a  kite,  359;  on  the  recogni- 
tion of  dogs  and  cats  by  ducks, 
406;  on  the  pairing  of  a  wild 
duck  with  a  pintail  drake,  409; 
on  the  courtship  of  fowls,  411; 
on  the  coupling  of  pheasants 
with  common  hens,  414. 

Hilgendorf,  sounds  produced  by 
crustaceans,  270. 

Hindoo,  his  horror  of  breaking 
his  caste,  118,  120. 

Hindoos,  local  difference  of  stat- 
ure among,  30;  difference  of, 
from  Europeans,  187;  color  of 
the  beard  in,  554. 

Hipparchia  Janira,  315;  instability 
of  the  ocellated  spots  of,  422. 


640 


INDEX; 


Hippocampus,  development  of, 
158;  marsupial  receptacles  of  the 
male,  342. 

minor,  198. 

Hippopotamus,    nakedness   of,   54. 

Hips,  proportions  of,  in  soldiers 
and  sailors,  31. 

Hodgson,  S.,  on  the  sense  of  duty, 
94. 

Hoffberg-,  on  the  horns  of  the 
reindeer,  499;  on  sexual  prefer- 
ences shown  by  reindeer,  520. 

Hoffman,  Prof.,  protective  colors, 
278;  fighting  of  frogs,  346. 

Hog-deer,  541. 

Hog,   wart-,  515;   river-,   516. 

Holland,  Sir  H.,  on  the  effects  of 
new  diseases,  178. 

Homologous  structures,  corre- 
lated variation  of,  42. 

Homoptera,  279;  stridulation  of 
the,   and  Orthoptera,  discussed, 

286. 

Honduras,  Quiscalus  major  m,  244. 

Honey-buzzard  of  India,  varia- 
tion in  the  crest  of,  418. 

Honey-suckers,  moulting  of  the, 
386;  Australian,  nidification  of, 
448. 

Honor,  law  of,  117. 

Hooker,  Dr.,  forbearance  of  ele- 
phant to  his  keeper,  101;  on  the 
color  of  the  beard  in  man,  553. 

Hookham,  Mr.,  on  mental  con- 
cepts in  animals,  81. 

Hoolock  Gibbon,  nose  of,  146. 

Hoopoe,  366;  sounds  produced  by 
the  male,  371. 

Hoplopterus  armatus,  wing-spurs 
of,  361. 

Hornbill,  African,  inflation  of  the 
neck-wattle  of  the  male  during 
courtship,   377. 

Hornbills,  sexual  difference  in  the 
color  of  the  eyes  in,  420;  nidifi- 
cation  and  incubation   of,   448. 

Home,  C,  on  the  rejection  of  a 
brightly-colored  locust  by  liz- 
ards and  birds,  287. 

Horns,  sexual  differences  of,  in 
sheep  and  goats,  225;  loss  of,  in 
female  merino  sheep,  226;  devel- 
opment of,  in  deer,  228;  develop- 
ment of,  in  antelopes,  229;  from 
the  head  and  thorax,  in  male 
beetles,  292;  of  deer,  499,  502,  510; 
and  canine  teeth,  inverse  devel- 
opment of,  510. 

Horse,  fossil,  extinction  of  the, 
in  South  America.  186;  polygam- 
ous, 213;  canine  teeth  of  male, 
498;  winter  change  of  color,  537. 

Horses,  rapid  increase  of,  in 
South  America,  45:  diminution 
of  canine  teeth  in,  52;  dreaming, 
72;  of  the  Falkland  Islands  and 
Pampas,  176;  numerical  propor- 
tion of  the  sexes  in,  211,  212: 
lighter  in  winter  in  Siberia,  224; 
sexual  preferences  in,  520;  pair' 


ing  preferently  with  those  of  the 
same  color,  535;  numerical  pro- 
portion of  male  and  female 
births  in,  240;  formerly  striped, 
542. 

Hottentot  women,  peculiarities  of, 
170. 

Hottentots,  lice  of,  166;  readily 
become  musicians,  565;  notions 
of  female  beauty  of  the,  573; 
compression  of  nose  by,   579. 

Hough,  Dr.  S.,  men's  temperature 
more  variable  than  v/omen's, 
220;  proportion  of  sexes  in  man, 
237. 

House-slaves,  difference  of,  from 
field-slaves,  191. 

Houzeau,  on  the  baying  of  the 
dog,  72;  on  reason  in  dogs,  74; 
birds  killed  by  telegraph  wires, 
78;  on  the  cries  of  domestic 
fowls  and  parrots,  82,  85;  ani- 
mals feel  no  pity,  99;  suicide  in 
the  Aleutian  islands,  113. 

Howorth,  H.  H.,  extinction  of 
savages,   179. 

Huber,  P.,  on  ants  playing  to- 
gether, 67;  on  memory  in  ants, 
72;  on  the  intercommunication 
of  ants,  87;  on  the  recognition 
of  each  other  by  ants  after  sep- 
aration, 289. 

Hue,  on  Chinese  opinions  of  the 
appearance  of  Europeans,  572. 

Huia,  the,  of  New  Zealand,  204. 

Human,  man  classed  alone  in  a, 
kingdom,  143. 

sacrifices,  93. 

Humanity,  unknown  among  some 
savages,  114;  deficiency  of, 
among  savages,  119. 

Humboldt,  A.  von,  on  the  ration- 
ality of  mules,  76;  on  a  parrot 
preserving  the  language  of  a 
lost  tribe,  177;  on  the  cosmetic 
arts  of  savages,  568;  on  the  ex- 
aggeration of  natural  characters 
by  man,  577;  on  the  red  painting 
of  American  Indians,  577. 

Hume,  D.,  on  sympathetic  feel- 
ings, 105. 

Humming-bird,  racket-shaped 

feathers  in  the  tail  of  a,  380; 
display  of  plumage  by  the  male, 
389. 

Humming-birds,  ornament  their 
nests,  89,  408;  polygamous,  214; 
proportion  of  the  sexes  in,  243, 
483;  sexual  differences  in,  356, 
436;  pugnacity  of  male,  356; 
modified  primaries  of  male,  373; 
coloration  of  the  sexes  of,  382; 
display  by,  437;  nidification  of 
the,  447;  colors  of  female,  447; 
young  of,  483. 

Humphreys,  H.  N.,  on  the  habits 
of  the   stickle-back,   216,  330. 

Hunger,    instinct   of,   108. 

Huns,  ancient,  flattening  of  the 
nose  by  the,  577. 


INDEX. 


641 


Hunter,  J.,  on  the  number  of  spe- 
cies of  man,  170;  on  secondary 
sexual  characters,  203;  on  the 
general  behavior  of  female  ani- 
mals during-  courtship,  217;  on 
the  muscles  of  the  larynx  in 
song--birds,  366;  on  the  curled 
frontal  hair  of  the  bull,  526;  on 
the  rejection  of  an  ass  by  a  fe- 
male zebra,  535. 

,     W.     W.,     on     the     recent 

rapid  Increase  of  the  Santali, 
44;  on  the  Santali,  187. 

Huss,  Dr.  Max,  on  mammary 
glands,  158. 

Hussey,  Mr.,  on  a  partridge  dis- 
tinguishing persons,   406. 

Hutchinson,  Col.,  example  of  rea- 
soning in  a  retriever,  75. 

Hutton,  Capt.,  on  the  male  wild 
goat  falling  on  his  horns,  503. 

Huxley,  T.  H.,  on  the  structural 
agreement  of  man  with  the 
apes,  2;  on  the  agreement  of  the 
brain  in  man  with  that  of  lower 
animals,  6;  on  the  adult  age  of 
the  orang,  8;  on  the  embryonic 
development  of  man,  9;  on  the 
origin  of  man,  2,  11;  on  varia- 
tion in  the  skulls  of  the  natives 
of  Australia,  25;  on  the  abductor 
of  the  fifth  metatarsal  in  apes, 
40;  on  the  nature  of  the  reason- 
ing power,  75;  on  the  position 
of  man,  146;  on  the  sub-orders 
of  primates,  148;  on  the  Lemu- 
rid^,  153;  on  the  Dinosauria, 
154;  on  the  amphibian  affinities 
of  the  Ichthyosaurians,  154;  on 
variability  of  the  skull  in  cer- 
tain races  of  man,  170;  on  the 
races  of  man,  172;  supplement 
on  the  brain,  199. 

Hybrid  birds,  production  of,  409. 

Hydrophobia  communicable  be- 
tween man  and  the  lower  ani- 
mals, 7. 

Hydroporus,  dimorphism  of  fe- 
males of,  273. 

Hyelaphus  porcinus,  541. 

Hygrogonus,   342. 

Hyla,   singing  species  of,  346. 

Hylobates,  absence  of  the  thumb 
in,  49;  upright  progression  of 
some  species  of,  51;  maternal  af- 
fection in  a,  68;  direction  of  the 
hair  on  the  arms  of  species  of, 
147;  females  of,  less  hairy  below 
than  males,   553. 

agilis,  49;  hair  on  the  arms 

of,  147;  musical  voice  of  the, 
522;  superciliary  ridge  of,  553; 
voice  of,  562. 

hoolock,  sexual  difference  of 


color  in,  532. 

lar,  49;  hair  on  the  arms  of, 


leuciscus,  49;  song  of,  562. 
syndactyius,   49;   laryng'e^l 


147. 


sac  of, 
41 


Hylophila   prasinana,    305. 

Hymenoptera,  289;  large  size  of 
the  cerebral  ganglia  in,  52;  class- 
ification of,  144;  sexual  differ- 
ences in  the  wings  of,  274;  acu- 
leate, relative  size  of  the  sexes 
of,  276. 

Hymenopteron,  parasitic,  with  a 
sedentary  male,  217. 

Hyomoschus  aquations,  442. 

Hyperythra,  proportion  of  the 
sexes  in,  246. 

Hypogymna  dispar,  sexual  differ- 
ence of  color  in,  312. 

Hypopyra,  coloration  of,  312. 


Ibex,  male,  falling  on  his  horns, 
503;   beard  of  the,   526. 

Ibis,  white,  change  of  color  of 
naked  skin  in,  during  the  breed- 
ing season,  384;  scarlet,  young  of 
the,  475. 

— tantalus,  age  of  mature  plu- 
mage in,  478;  breeding  in  imma- 
ture plumage,  479. 

Ibises,  decomposed  feathers  in, 
380;  white,  488;  and  black,  489. 

Ichneumonidae,  difference  of  the 
sexes  in,    289. 

Ichthyopterygia,  36. 

Ichthyosaurians,  154. 

Idiots,  microcephalous,  their 
characters  and  habits,  34;  hairi- 
ness and  animal  nature  of  their 
actions,  35;  microcephalous,  im- 
itative faculties  of,  85. 

Iguana  tuberculata,  350. 

Iguanas,  350. 

Illegitimate  and  legitimate  chil- 
dren, proportion  of  the  sexes  in, 
239. 

Imagination,  existence  of,  in  ani- 
mals, 72. 

Imitation,  66;  of  man  by  mon- 
keys, 71;  tendency  to,  in  mon- 
keys, microcephalous  idiots  and 
savages,  85;   influence  of,  126. 

Immature  plumage  of  birds,  456, 
459. 

Implacentata,  153. 

Implements,  employed  by  mon- 
keys, 78;  fashioning  of,  peculiar 
to  man,  80. 

Impregnation,  period  of,  influence 
of,    upon  sex,   239. 

Improvement,  progressive,  man 
alone  supposed  to  be  capable  of, 
77. 

Incisor  teeth,  knocked  out  or  filed 
by  some  savages,  569. 

Increase,  rate  of,  43;  necessity  of 
checks   in,    45. 

Indecency,  hatred  of,  a  modern 
virtue,    115. 

India,  difficulty  of  distinguishing 
the  native  races  of,  163;  Cyprin- 
idse  of,  S99;  color  of  the  bfeard 
in  ratfes  tff  men  of,  553, 


642 


INDEX. 


Indian,  North  American,  honored 
for  scalping-  a  man  of  another 
tribe,  113. 

Individuality,   in  animals,  81. 

Indopicus  carlotta,  colors  of  the 
sexes  of,  452. 

Infanticide,  prevalence  of,  45,  113, 
253;  s;apposed  cause  of,  572;  pre- 
valence and  causes  of,  586  et 
seq. 

Inferiority,  supposed  physical,  of 
man,   61. 

Inflammation  of  the  >  bowels,  oc- 
currence of,  in  Cebus  Azarse,  7. 

Inheritance,  27;  of  long  and  short 
sig-ht,  32;  of  effects  of  use  of 
vocal  and  mental  organs,  86;  of 
moral  tendencies,  119,  122;  laws 
of,  222;  sexual,  227;  sexually  lim- 
ited, 439. 

Inquisition,   influence   of  the,    138. 

Insanity,  hereditary,  27. 

Insect,  fossil,  from  the  Devonian, 
286. 

Insectivora,  529;  absence  of  sec- 
ondary sexual  characters  in,  214. 

Insects,  relative  size  of  the  cere- 
bral ganglia  in,  52;  male,  ap- 
pearance of,  before  the  females, 
208;  pursuit  of  female,  by  the 
males,  217;  period  of  develop- 
ment of  sexual  characters  in, 
231;  secondary  sexual  characters 
of,  272;  stridulation,  561. 

Insessores,  vocal  organs  of,  366. 

Instep,  depth  of,  in  soldiers  and 
sailors,  31. 

Instinct  and  intelligence,  65. 

,  migratory,  vanquishing  the 

maternal,  104,  109. 

Instinctive  actions,  the  result  of 
inheritance,  102. 

impulses,  difference  of    the 

force  of,  107;  and  moral  im- 
pulses,  alliance  of,   107. 

Instincts,  64;  complex  origin  of, 
through  natural  selection,  65; 
possible  origin  of  some,  65;  ac- 
quired, of  domestic  animals,  101; 
variability  of  the  force  of,  104; 
difference  of  force  between  the 
social  and  other,  108,  122;  utilized 
for  new  purposes,  565. 

Instrumental  music  of  birds,  371, 
374. 

Intellect,  influence  of,  in  natural 
selection  in  civilized  society,  133. 

Intellectual  faculties,  their  influ- 
ence on  natural  selection  in 
man,  124;  probably  perfected 
through  natural  selection,  125. 

Intelligence,  Mr.  H.  Spencer  on 
the  dawn  of,  65. 

Intemperance,  no  reproach  among 
savages,  116;  its  destructive- 
ness,  134. 

Intoxication  in  monkeys,  7. 

Iphias   glaucippe,   309. 

Iris,  sexual  difference  in  the  color 
ot  thte,  in  birds,  377,  419. 


Ischio-pubic  muscle,  40. 
Ithaginis     cruentus,     number     of 

spurs  in,  360. 
lulus,  tarsal  suckers  of  the  males 

of,  271. 


Jackals,  learning  from  dogs  to 
bark,  70. 

Jack-snipe,  coloration  of  the,  486. 

Jacquinot,  on  the  number  of  spe- 
cies of  man,  170, 

Jaeger,  Dr.,  length  of  bones  in- 
creased from  carrying  weights, 
31;  on  the  diflficulty  of  approach- 
ing herds  of  wild  animals,  97; 
male  Silver-pheasant,  rejected 
when  his  plumage  was  spoilt, 
413. 

Jaguars,  black,  535. 

Janson,  E.  W.,  on  the  proportions 
of  the  sexes  in  Tomicus  villosus, 
250;    on  stridulant  beetles,  299. 

Japan,  encouragement  of  licen- 
tiousness in,  45. 

Japanese,  general  beardlessness  of 
the,  555;  aversion  of  the,  to 
whiskers,  275. 

Jardine,  Sir  W.,  on  the  Argus 
pheasant,  379,  397. 

Jarrold,  Dr.,  on  modifications  of 
the  skull  induced  by  unnatural 
position,  54. 

Jarves,  Mr.,  on  infanticide  in  the 
Sandwich  Islands,  254. 

Javans,  relative  height  of  the 
sexes  of,  554;  notions  of  female 
beauty,   574. 

Jaw,  influence  of  the  muscles  of 
the,  upon  the  physiognomy  of 
the  apes,   52. 

Jaws,  smaller  proportionately  to 
the  extremities,  32;  influence  of 
food  upon  the  size  of,  32;  dimin- 
ution of,  in  man,  51;  in  man,  re- 
duced by  correlation,  557. 

Jay,  young  of  the,  475;  Canada, 
young  of  the,  475. 

Jays,  new  mates  found  by,  402; 
distinguishing  persons,  406. 

Jeffreys,  J.  Gwyn,  on  the  form  of 
the  shell  in  the  sexes  of  the 
Gasteropoda,  259;  on  the  influ- 
ence of  light  upon  the  colors  of 
shells,    260. 

Jelly-fish,  bright  colors  of  some, 
257. 

Jenner,  Dr.,  on  the  voice  of  the 
rook,  370;  on  the  finding  of  new 
mates  by  magpies,  402;  on  re- 
tardation of  the  generative 
functions  in  birds.  404. 

Jenyns,  L.,  on  the  desertion  of 
their  young  by  swallows,  104;  on 
male  birds  singing  after  the 
proper  season,  404. 

Jferdon,  Dr.,  on  birdg  dreaming. 
72;  on  thfe  prusnacity  of  thfe  male 


INDEX. 


643 


bulbul,  357;  on  the  pugnacity  of 
the  male  Ortygornis  gularis, 
359;  on  the  spurs  of  Galloperdix, 
360;  on  the  habits  of  Lobivanel- 
lus,  362;  on  the  spoonbill,  370;  on 
the  drumming  of  the  Kalij- 
pheasant,  371;  on  Indian  bus- 
tards, 373;  on  Otis  bengalensis, 
376;  on  the  ear-tufts  of  Sypheo- 
tides  auritus,  379;  on  the  double 
moults  of  certain  birds,  386;  on 
the  moulting  of  the  honej'-- 
svickers,  386;  on  the  moulting  of 
bustards,  plovers,  and  drongos, 
387;  on  the  spring  change  of 
color  in  some  finches,  388;  on 
display  in  male  birds,  389;  on  the 
display  of  the  under-tail  coverts 
by  the  male  bulbul,  396;  on  the 
Indian  honey-buzzard,  418;  on 
sexual  differences  in  the  color 
of  the  eyes  of  horn-bills,  420;  on 
the  markings  of  the  Tragopan 
pheasant,  422;  on  the  nidification 
of  the  Orioles,  447;  on  the  nidi- 
fication of  the  hornbills,  448;  on 
the  Sultan  yellow-tit,  451;  on 
Palseornis  javanicus,  455;  on  the 
imm.ature  plumage  of  birds,  459 
et  seq. ;  on  representative  spe- 
cies of  birds,  462;  on  the  habits 
of  Turnix,  470;  on  the  continued 
increase  of  beauty  of  the  pea- 
cock, 480;  on  coloration  in  the 
genus  Palieornis,  489. 

Jevons,  W.  S.,  on  the  migrations 
of  man,   46. 

Jews,  ancient  use  of  flint  tools 
by  the,  141;  uniformity  of,  in 
various  parts  of  the  world,  1S8: 
numerical  proportion  of  male 
and  female  births  among  the, 
237;  ancient,  tattooing  practiced 
by,   569. 

Johnstone,  Lieut.,  on  the  Indian 
elephant,  213. 

Jollofs,  fine  appearance  of  the, 
582. 

Jones,  Albert,  proportion  of  sexes 
of  Lepidoptera,  reared  by,  249. 

Juan  Fernandez,  humming-birds 
of,  483. 

Junonia,  sexual  differences  of  col- 
oring in  species  of,  307. 

Jupiter,  comparison  with  Assyri- 
an effigies,  576. 


K 

Kaffir  skull,  occurrence  of  the 
diastema  in  a,  39. 

Kaffirs,  their  cruelty  to  animals, 
114;  lice  of  the,  166;  color  of  the, 
574;  engrossment  of  the  hand- 
somest women  by  the  chiefs  of 
the,  5S0;  marriage-customs  of 
tlie   593. 

_Kalij-ph«asant,  drumming  o'f  the 
mm%  373' ;  ypMlxg  of,  463. 


Kallima,  resemblance  of,  to  a 
withered  leaf,  308. 

Kalmucks,  general  beardlessness 
of,  555;  aversion  of,  to  hairs  on 
the  face,  575;  marriage-customs 
of  the,  592. 

Kangaroo,  great  red,  sexual  dif- 
ference in  the  color  of,  529. 

Kant,  Imm.,  on  duty,  94;  on  self- 
restraint,  106;  on  the  number  of 
species  of  man,  170. 

Katy-did,  stridulation  of  the,  280. 

Keen,  Dr.,  on  the  mental  powers 
of  snakes,  348. 

Keller,  Dr.,  on  the  difficulty  of 
fashioning  stone  implements,  48. 

Kent,  W.  S.,  elongation  of  dorsal 
fin  of  Callionymus  lyra,  333; 
courtship  of  Labrus  mixtus,  337; 
colors  and  courtship  of  Can- 
tharus  lineatus,   337. 

Kestrels,  new  mates  found  by, 
402. 

Kidney,  one,  doing  double  work 
in  disease,  31. 

King,  W.  R.,  on  the  vocal  organs 
of  Tetrao  cupido,  368;  on  the 
drumming  of  grouse,  371;  on  the 
reindeer,  499;  on  the  attraction 
of  male  deer  by  the  voice  of  the 
female,  522. 

King  and  Fitzroy,  on  the  mar- 
riage-customs of  the  Fuegians, 
593. 

King-crows,  nidification  of,  447. 

Kingfisher,  366;  racket-shaped 
feathers  in  the  tail  of  a,  379. 

Kingfishers,  colors  and  nidifi.ca- 
tion  of  the,  449,  451,  452;  imma- 
ture plumage  of  the,  461,  463; 
young  of  the,  475. 

King  Lory,  451;  immature  plu- 
mage of  the,  461. 

Kingsley,  C,  on  the  sounds  pro- 
duced by  Umbrina,  343. 

Kirby  and  Spence,  on  sexual  dif- 
ferences in  the.  length  of  the 
snout  in  Curculionidse^  204;  on 
the  courtship  of  insects.  217;  on 
the  elytra  of  D^tiscus,  274;  on 
peculiarities  in  the  legs  of  male 
insects,  274;  on  the  relative  size 
of  the  sexes  in  insects.  276;  on 
the  Fulgoridas,  279;  on  the  hab- 
its of  Termites,  288;  on  differ- 
ence of  color  in  the  sexes  o'f  bee- 
tles, 291;  on  the  horns  of  the 
male  lamellicorn  beetles,  294;  on 
hornlike  processes  in  male  Cur- 
culionidse,  296;  on  the  pugnacity 
of  the  male  stag-beetle,  297. 

Kite,  killed  by  a  game-cock,  359. 

Knot,  retention  of  winter  plu- 
mage by  the,  386. 

Knox,  R.,  on  the  semilunar  fold, 
17;  on  the  occurrence  of  the 
supra.-condyloid  foramen  in  the 
humefrus  of  naan,  21;  on  tire  fea- 
ttfre's  of  "tm  yoSins  Mfe^mntJn,  164. 


644 


INDEX. 


Koala,  length  of  the  Cjccum   in, 

20. 
Kobus  ellipsiprymnus,  proportion 

of  the  sexes  in,  242. 
Kolreuter,  on  the  sterility  of  hy- 
brid plants,   168. 
Koodoo,  development  of  the  horns 

of  the,  229;  markings  of  the,  538. 
Koppen,   F.  T.,  on  the  migratory 

locust,  280. 
Kordofan,     protuberances     artifi- 
cially produced    by    natives  of, 

569. 
Koraks,  marriage  customs  of,  593. 
Korte,  on  the  proportion  of  sexes 

in  locusts,  250;   Russian  locusts, 

280. 
Kovalevsky,  A.,  on  the  affinity  of 

the   Ascidia  to   the  Vertebrata, 

155. 
,  W.,  on  the  pugnacity  of  the 

male    capercailzie,    359;    on    the 

pairing  of  the  capercailzie,  362. 
Krause,   on  a  convoluted  body  at 

the   extremity  of  the   tail   in   a 

Macacus  and  a  cat,  22. 
Kupffer,   Prof.,  on  the  affinity  of 

the  Ascidia  to  the  Vertebrata, 

155. 


Labidocera    Darwinii,    prehensile 
organs  of  the  male,  263. 

Labrus,  splendid  colors  of  the  spe- 
cies of,  338. 

mixtus,     sexual  differences 


in,    334,    337. 
pavo,  339. 


Lacertilia,  sexual  differences  of, 
350. 

Lafresnaye,  M.  de,  on  birds  of 
paradise,   381. 

Lamarck,  on  the  origin  of  man,  2. 

Lamellibranchiata,  259. 

Lamellicorn  beetles,  horn-like 
processes  from  the  head  and 
thorax  of,  292,  296;  influence  of 
sexual   selection  on,  298. 

Lamellicornia,  stridulation  of,  300. 

Lamont,  Mr.,  on  the  tusks  of  the 
walrus,  498;  on  the  use  of  its 
tusks  by  the  walrus,  509;  on  the 
bladder-nose  seal,  524. 

Lampornis  porphyrurus,  colors  of 
the  female.  447. 

Lampyridas,  distasteful  to  mam- 
mals, 475. 

Lancelet,  154,  160. 

Landois,  H.,  gnats  attracted  by 
sound,  278;  on  the  production  of 
sound  by  the  Cicadse,  279;  on 
the  stridulating  organ  of  the 
crickets,  281;  on  Decticus,  283; 
g_n  the  stridulating  organs,  of 
the  Acridiidfe,  284;  stridulating 
apparatus  in  Orthoptera,  286; 
sounds  produced  by  Atropus, 
289;     on.    th^     sttiQiila,13<cm     ot 


Necrophorus,  299;  on  the 
stridulant  organ  of  Cerambyx 
heros,  300;  on  the  stridulant 
organ  of  Geotrupes,  300;  on  the 
stridulating  organs  in  the  Cole- 
optera,  301;  on  the  ticking  of 
Anobium,  303. 

Landor,  Dr.,  on  remorse  for  not 
obeying   tribal   custom.    111. 

Language  an  art,  83;  articulate, 
origin  of,  84;  relation  of  the 
progress  of,  to  the  development 
of  the  brain,  85j  effects  of  in- 
heritance in  production  of,  86; 
complex  structure  of,  among 
barbarous  nations,  88;  natural 
selection  in,  88;  gesture,  174; 
primeval,  176;  of  a  lost  tribe  pre- 
served by  a  parrot,  177. 

Languages,  presence  of  rudi- 
ments in,  88;  classification  of,  88; 
variability  of,  88;  crossing  or 
blending  of,  88;  complexity  of, 
no  test  of  perfection  or  proof  of 
special  creation,  89;  resemblance 
of,  evidence  of  community  of 
origin,  144. 

and  species,  identity  a£  evi- 
dence of  their  gradual  develop- 
ment,  87. 

Lanius,  455;  characters  of  young, 
459. 

rufus,  anomalous  young    of 

477. 

Lankester,  E.  R.,  on  comparative 
longevity,  130,  132;  on  the  de- 
structive effects  of  intemper- 
ance, 134. 

Lanugo,  of  the  human  foetus,  19, 
594. 

Lapponian  language,  highly  arti- 
ficial, 88. 

Lark,  proportion  of  the  sexes  in 
the,  243;  female,  singing  of  the, 
365. 

Larks,  attracted  by  a  mirror,  407. 

Lartet,  E.,  comparison  of  cranial 
capacities  of  skulls  of  recent 
and  tertiary  mammals,  53;  on 
the  size  of  the  brain  in  mam- 
mf!ils,  78;  on  Dryopithecus,  151; 
en  pre-historic  flutes,  565. 

Larus,  seasonal  change  of  plu- 
mage in,  488. 

Larva,  luminous,  of  a  Brazilian 
beetle,  275. 

Larynx,  muscles  of  the,  in  song- 
birds, 366. 

Lasiocampa  quercus,  attraction  of 
males  by  the  female,  248;  sexual 
difference  of  color  in,  312. 

Latham,  R.  G.,  on  the  migrations 
of  man,  46. 

Latooka,  perforation  of  the  lower 
lip  by  the  women  of,  570. 

Laurillard,  on  the  abnormal  di- 
vision of  the  malar  bone  in  man, 
37. 

Lawrence,  W.,  on  thp  superiority 
o'f  sa.vagefs  tb  i^urb'pteians  in  p'qV" 


INDEX. 


645 


er  of  sight,  32;  on  the  color  of 
negro  infants,  553;  on  the  fond- 
ness of  savages  for  ornaments, 
568;  on  beardless  races,  575;  on 
the  beauty  of  the  English  aristo- 
cracy, 581. 

Layard,  E.  L.,  on  an  instance  of 
rationality  in  a  cobra,  348;  on 
the  pugnacity  of  Gallus  Stan- 
leyi,   359. 

Laycock,  Dr.,  on  vital  periodicity, 
8;  theroid  nature  of  idiots,  35. 

Leaves,  autumn,  tints  useless, 
259. 

Lecky,  Mr.,  on  the  sense  of  duty, 
94;  on  suicide,  113;  on  the  prac- 
tice of  celibacy,  115;  his  view  of 
the  crimes  of  savages,  116;  on 
the  gradual  rise  of  morality,  121. 

Leconte,  J.  L.,  on  the  stridulant 
organ  in  the  Coprini  and  Dy- 
nastini,  300. 

Lee,  H.,  on  the  numerical  pro- 
portion of  the  sexes  in  the  trout, 
245. 

Leg,  calf  of  the.  artificially  mod- 
ified, 569. 

Legitimate  and  illegitimate  chil- 
dren, proportion  of  the  sexes  in, 
239. 

Legs,  variation  of  the  length  of 
the,  in  man,  25;  proportions  of, 
in  soldiers  and  sailors,  31;  front, 
atrophied  in  some  male  butter- 
flies, 274;  peculiarities  of,  in 
male  insects,  275. 

Leguay,  on  the  occurrence  of  the 
inter-condyloid  foramen  in  the 
humerus  of  man,  21. 

"Lek"  of  the  black-cock  and  cap- 
ercailzie, 389. 

Lemoine,  Albert,  on  the  origin  of 
language,   84. 

Lemur  macaco,  sexual  difference 
of  color  in,  532. 

Iiemuridse,  148;  ears  of  the,  15; 
variability  of  the  muscles  in  the, 
40;  position  and  derivation  of 
the,  153;  their  origin,  161. 

Lemurs,  uterus  in  the,   37. 

Lenguas,  disfigurement  of  the 
ears  of  the,  570. 

Leopards,  black,  535. 

Lepidoptera,  304;  numerical  pro- 
portions of  the  sexes  in  the,  245; 
coloring  of,  305;  ocellated  spots 
of,   422. 

Lepidosiren,  154,  160. 

Leptalides,  mimicry  of,  321. 

Leptorhynchus  angustatus,  pug- 
nacity of  male,  299. 

Leptura  testacea,  difference  of 
color  in  the  sexes  of,  296. 

Leroy,  on  the  wariness  of  young 
foxes  in  hunting-districts,  78;  on 
the  desertion  of  their  young  by 
swallows,  104. 

Ibeslie,  D.,  marriage-customs  of 
Kaffirs,  ^3. 

Lesse,  valles^  o^f  the,  21. 
42 


Lesson,  on  the  birds  of  paradise, 
214,  397;  on  the  sea-elephant,  524, 

Lessona,  M.,  observations  on 
Serranus,  157. 

Lethrus  cephalotes,  pugnacity  of 
the  males  of,  294,  297. 

Leuckart,  R.,  on  the  vesicula 
prostatica,  23;  on  the  influence 
of  the  age  of  parents  on  the  sex 
of  offspring,  239. 

Levator  claviculee  muscle,  40. 

Libellula  depressa,  color  of  the 
male,  28S. 

Libellulidse,  relative  size  of  the 
sexes  of,  276;  difference  in  the 
sexes  of,  287. 

Lice  of  domestic  animals  and 
man,    165. 

Licentiousness,  a  check  upon  pop- 
ulation, 45;  prevalence  of, 
among  savages,  115. 

Lichtenstein,  on  Chera  progne, 
413. 

Life,  inheritance  at  corresponding 
periods  of,  223,  227. 

Light,  effects  on  complexion,  31; 
influence  of,  upon  the  colors  of 
shells,  260. 

Lilford,  Lord,  the  ruff  attracted 
by  bright  objects,  407. 

Limosa  lapponica,  472. 

Linaria,  455. 

montana,  243. 

Lindsay,  Dr.  W.  L.,  diseases 
communicated  from  animals  to 
man,  7;  madness  in  animals,  77; 
the  dog  considers  his  master  his 
God,  93. 

Linnaeus,  views  of,  as  to  the  po- 
sition of  man,  145~. 

Linnet,  numerical  proportion  of 
the  sexes  in  the,  243;  crimson 
forehead  and  breast  of  the,  388; 
courtship  of  the,  395. 

Lion,  polygamous,  214;  mane  of 
the,  defensive,  516;  roaring  of 
the,  522. 

Lions,    stripes   of   young,   458. 

Lips,  piercing  of  the,  by  savages, 
570. 

Lithobius,  prehensile  appendages 
of  the  female,   271. 

Lithosia,  coloration  in,  311. 

Littorina  littorea,  259. 

Livingstone,  Dr.,  manner  of  sit- 
ting of  gorilla,  147;  on  the  in- 
fluence of  dampness  and  dry- 
ness on  the  color  of  the  skin, 
188;  on  the  liability  of  negroes 
to  tropical  fevers  after  residence 
in  a  cold  climate,  189;  on  the 
spur-winged  goose,  360;  on  wea- 
ver-birds, 371;  on  an  African 
night-jar,  379,  397;  on  the  bat- 
tle-scars of  South  African  male 
mammals,  496;  on  the  removal 
of  the  upper  incisors  by  the 
Batokas,  569;  on  the  perforation 
of  the  upper  lip  by  the  Maka« 
ItjlDi  {JTOj  xm  the  Daafyai,  574. 


646 


INDEX. 


Livonia,  numerical  proportion  of 
male  and  female  births  in,  211, 
238. 

Lizards,  relative  size  of  the  sexes 
of,   350;   g-ular  pouches   of,   350. 

Lloyd,  L.,  on  the  polygamy  of  the 
capercailzie  and  bustard,  215;  on 
the  numerical  proportion  of  the 
sexes  in  the  capercailzie  and 
black-cock,  242;  on  the  salmon, 
330;  on  the  colors  of  the  sea 
scorpion,  334;  on  the  pugnacity 
of  male  grouse,  360;  on  the  ca- 
percailzie and  black-cock,  362, 
366;  on  the  call  of  the  capercail- 
zie, 370;  on  assemblages  of 
grouse  and  snipes,  400,  on  the 
pairing  of  a  shield-drake  with 
a  common  duck,  409;  on  the  bat- 
tles of  seals,  496;  on  the  elk,  503. 

Lobivanellus,  wing-spurs  in,  361. 

Local  influences,  effect  of,  upon 
stature,  30. 

Lockwood,  Mr.,  on  the  develop- 
ment of  Hippocampus,  158. 

,  Rev.  S.,  musical  mouse,  562. 

Locust,  bright-colored,  rejected  by 
lizards  and  birds,  287. 

,  migratorj',  280;  selection  by 

female,  281, 

Locusts,  proportion  of  sexes  in, 
250;   stridulation  of,  281. 

Locustidse,  stridulation  of  the, 
280,  282;  descent  of  the,  282. 

Longicorn  beetles,  difference  of 
the  sexes  of,  in  color,  291;  strid- 
ulation of,   300. 

Lonsdale,  Mr.,  on  an  example  of 
personal  attachment  in  Helix 
pomatia,   260. 

Lophobranchii,  marsupial  recept- 
acles of  the  males,   342. 

Lophophorus,  habits  of,  414. 

Lophorina  atra,  sexual  difference 
in  coloration  of,   487. 

Lophornis  ornatus,  382. 

Lord,  J.  K.,  on  Salmo  lyeaodon, 
331. 

Lory,  King,  451;  immature  plu- 
mage of  the,  461. 

Love-antics  and  dances  of  birds, 
375. 

Lowne,  B.  T.,  on  Musea  vomito- 
ria,    52,    278. 

Loxia,  characters  of  young  of,  459. 

Lubbock,  Sir  J.,  on  the  antiquity 
of  man,  2;  on  the  origin  of  man, 
2;  on  the  mental  capacity  of 
savages,  63;  on  the  origin  of 
implements,  80;  on  the  simplifi- 
cation of  languages,  89;  on  the 
absence  of  the  idea  of  God 
among  certain  races  of  men,  91; 
on  the  origin  of  the  belief  in 
spiritual  agencies,  91;  on  super- 
stitions, 94;  on  the  sense  of  duty, 
94;  on  the  practice  of  burying 
the  did  and  sitik  among  thie  Fi- 


jians,  99;  on  the  immorality  of 
savages,  116;  on  Mr.  Wallace's 
claim  to  the  origination  of  the 
idea  of  natural  selection,  47;  on 
the  absence  of  remorse  among 
savages,  128;  on  the  former  bar- 
barism of  civilized  nations,  140; 
on  improvements  in  the  arts 
among  savages,  141;  on  resem- 
blances of  the  mental  charac- 
ters in  different  races  of  men, 
174;  on  the  arts  practiced  by 
savages,  175;  on  the  power  of 
counting-  in  primeval  man,  175; 
on  the  prehensile  organs  of  the 
male  Labidocera  Darwinii,  263; 
on  Chlceon,  272;  on  Smynthurus 
luteus,  277;  "finding  of  new  mates 
by  jays,  402;  on  strife  for 
women  among  the  North  Ameri- 
can Indians,  556;  on  music,  565; 
on  the  ornamental  practices  of 
savages,  568;  on  the  estimation 
of  the  beard  among  the  Anglo- 
Saxons,  575;  on  artificial  deform- 
ation of  the  skull,  577;  on  "com- 
munal marriages,"  582,  583;  on 
exogamy,  584,  587;  on  the  Ved- 
dahs,  591;  on  polyandry,  587. 

liucanidae,  variability  of  the 
mandibles  in  the  male,  297. 

Lucanus,  large  size  of  males  of, 
276. 

cervus,  numerical  propor- 
tion of  sexes  of,  249;  weapons  of 
the  male,  297. 

elaphus,    tise    of   mandibles 


of,  298;  large  jaws  of  male,  273. 
Lucas,    Prosper,    on   pigeons,    412; 

on   sexual  preference   in  horses 

and  bulls,  520. 
Lunar  periods,  8,  156. 
Lund,    Dr.,    on    skulls    found     in 

Brazilian  caves,  164. 

Lungs,  enlargement  of,  in  the 
Quichua  and  Aymara  Indians, 
32;  a  modified  swim-bladder,  156; 
different  capacity  of  in  races  of 
man,  163. 

Luminosity  in  insects,  275. 

Luschka,  Prof.,  on  the  termina- 
tion of  the  coccyx,  22. 

Luxury,  expectation  of  life  unin- 
fluenced by,   132. 

Lycsena,  sexual  differences  of 
coloring  in  species  of,  307. 

Lyell,  Sir  C,  on  the  antiquity  of 
man,  2;  on  the  origin  of  man, 
2;  on  the  parallelism  of  the  de- 
velopment of  species  and  lan- 
guages, 87;  on  the  extinction  of 
languages,  88;  on  the  Inquisi- 
tion, 138;  on  the  fossil  remains 
of  vertebrata,  152;  on  the  fertil- 
ity of  mulattoes,  167. 

Lynx,  Canadian,  throat-ruff  of 
the,  516. 

Lyre^Td,'  assBHaMies  of,  401. 


INDBX. 


647 


M 


Macacus,  ears  of,  15;  convoluted 
body  in  the  extremity  of  the 
tail  of,  22;  variability  of  the  tail 
in  species  of,  56;  whiskers  of 
species  of,  527. 

brunneus,  57. 

cynomolgus,       superciliary 

ridge  of,  553;  beard  and  whiskers 
of,  becoming  white  with  age, 
554. 

ecaudatus,  5S. 

lasiotus,  facial  spots  of,  546. 

radiatus,  146. 

rhesus,  sexual  difference  in 

the  color  of,  534,  546. 

Macalister,  Prof.,  on  variations  of 
the  palmaris  accessorius  muscle, 
26;  on  muscular  abnormalities  in 
man,  40,  41;  on  the  greater  va- 
riability of  the  muscles  in  men 
than  in  women,  219. 

Macaws,  Mr.  Buxton's  observa- 
tions on,   99;   screams  of,  370. 

McCann,  J.,  on  mental  individual- 
ity, 82. 

McClelland,  J.,  on  the  Indian  Cy- 
prinid£e,   339. 

Macculloch,  Col.,  on  an  Indian  vil- 
lage without  any  female  chil- 
dren, 587. 

,    Dr.,    on  tertian  ague  in   a 

dog,  8. 

Macgillivray,  W.,  on  the  vocal 
organs  of  birds,  87;  on  the 
Egyptian  goose,  360;  on  the  hab- 
its of  wood-peckers,  371;  on  the 
habits  of  the  snipe,  373;  on  the 
whitethroat,  376;  on  the  moult- 
ing of  the  snipes,  386;  on  the 
moulting  of  the  Anatidee,  388;  on 
the  finding  of  new  mates  by 
magpies,  402;  on  the  pairing  of  a 
blackbird  and  thrush,  409;  on 
pied  ravens,  418;  on  the  guille- 
mots, 418;  on  the  colors  of  the 
tits,  452;  on  the  immature  plu- 
mage of  birds,  460  et  seq. 

Machetes,  sexes  and  young  of, 
480. 

pugnax,     supposed     to     be 

polygamous,  215;  numerical  pro- 
portion of  the  sexes  in,  243;  pug- 
nacity of  the  male,  357;  double 
moult  in,  385. 

Mcintosh,  Dr.,  colors  of  the  Ne- 
mertians,  261. 

McKennan,  marriage  customs  of 
Koraks,   593. 

Mackintosh,  on  the  moral  sense, 
94. 

MacLachlan,  R.,  on  Apatania 
muliebris  and  Boreus  hyemalis, 
251;  on  the  anal  appendages  of 
male  insects,  272;  on  the  pairing 
of  dragon-flies,  £76;  on  dragon- 
flies,  288;  on  dimorphism  in  Ag- 
rion,  2S8;  on  the  wa;nt  of  pug- 
nacity in  male  d'raSon-faes,  28S; 


color  of  ghost-moth  in  the  Shet- 
land Islands,   Zlt 

McLennan,  Mr.,  on  infanticide, 
45,  586;  on  the  origin  of  the  belief 
in  spiritual  agencies,  91;  on  the 
prevalence  of  licentiousness 
among  savages,  115,  583;  on  the 
primitive  barbarism  of  civilized 
nations,  140;  on  traces  of  the 
custom  of  the  forcible  capture 
of  wives,  140,  587;  on  polyandry, 
587. 

Macnamara,  Mr.,  susceptibility  of 
Andaman  islanders  and  Nepa- 
lese  to  change,  183. 

McNeill,  Mr.,  on  the  use  of  the 
antlers  of  deer,  506;  on  the 
Scotch  deerhound,  512;  on  the 
long  hairs  on  the  throat  of  the 
stag,  517;  on  the  bellowing  of 
stags,   521. 

Macropus,  courtship  of,  337. 

Macrorhinus  proboscideus,  struc- 
ture of  the  nose  of,  523. 

Magpie,  power  of  speech  of,  87; 
nuptial  assemblies  of,  401:  new 
mates  found  by,  402;  stealing 
bright  objects,  408;  young  of  the, 
475;   coloration  of  the,  489. 

Magpies,  vocal  organs  of  the,  366. 

Maillard,  M.,  on  the  proportion  of 
the  sexes  in  a  species  of  Papilio 
from  Bourbon,  246. 

Maine,  Sir  Henry,  on  the  absorp- 
tion of  one  tribe  by  another,  125; 
a  desire  for  improvement  not 
general,   129. 

Major,  Dr.  C.  Forsyth,  on  fossil 
Italian  apes,  151;  skull  of  Bos 
etruscus,  501;  tusks  of  miocene 
pigs,  516. 

Makalolo,  perforation  of  the  up- 
per lip  by  the,  570. 

Malar  bone,  abnormal  division  of, 
in  man,  37. 

Malay,  Archipelago,  marriage- 
customs  of  the  savages  of  the, 
593. 

Malays,  line  of  separation  be- 
tween the  Papuans  and  the,  165; 
general  beardlessness  of  the, 
555;  staining  of  the  teeth  among, 
569;  aversion  of  some,  to  hairs 
on  the  face,  575. 

— and    Papuans,     contrasted 

characters  of,  164. 

Male  animals,  struggles  of,  for 
the  possession  of  the  females, 
210;  eagerness  of,  in  courtship, 
216,  217;  generally  more  modi- 
fied than  female,  216,  218;  differ 
in  the  same  way  from  females 
and  young,  227. 

characters,  developed  in  fe- 
males, 223;  transfer  of,  to  female 
birds,  465. 

-,  sedentary,   of  a  hymenog* 


terous  parasite^  317. 
Maite€!abWrs,  ISS. 


64S 


INDEX. 


Males,  presence  of  rudimentary 
female  organs  in,  157, 

and    females,     comparative 

numbers  of,  209,  211;  compara- 
tive mortality  of,  while  young-, 
211. 

Malherbe,  on  the  woodpeckers, 
452. 

Mallotus  peronii,  328. 

villosus,  327. 

Malthus,  T.,  on  the  rate  of  in- 
crease of  population,  43,  44,  45. 

Maluridse,   nidification  of  the,  448. 

Malurus,  young  of,  480. 

Mammae,  204;  rudimentary,  in 
male  mammals,  11,  23,  157,  158; 
supernumerary,  in  women,  35;  of 
male  human  subject,  35. 

Mammalia,  Prof.  Owen's  classifi- 
cation of,  144;  genealogy  of  the, 
153. 

Mammals,  recent  and  tertiary, 
comparison  of  cranial  capacity 
of,  53;  nipples  of,  157;  pursuit  of 
female,  by  the  m.ales,  216;  sec- 
ondary sexual  characters  of,  496; 
weapons  of,  497;  relative  size  of 
the  sexes  of,  511;  parallelism  of, 
with  birds  in  secondary  sexual 
characters,  536;  voices  of,  used 
especially  during  the  breeding 
season,  562. 

Man,  variability  of,  25;  erroneous- 
ly regarded  as  more  domesti- 
cated than  other  animals,  27; 
migrations  of,  46;  wide  distribu- 
tion of,  46;  causes  of  the  naked- 
ness of,  55;  supposed  physical  in- 
feriority of,  61;  a  member  of  the 
Catarrhine  group,  151;  early 
progenitors  of,  156;  transition 
from  ape  indefinite,  176;  numer- 
ical proportions  of  the  sexes  in, 
210;  difference  between  the  sexes, 
218;  proportion  of  sexes  amongst 
the  illegitimate,  239;  different 
complexion  of  male  and  female 
negroes,  551;  secondary  sexual 
characters  of,  551;  primeval 
condition   of,    588. 

Mandans,  correlation  of  color  and 
texture  of  hair  in  the,  192. 

Mandible,  left,  enlarged  in  the 
male  of  Taphroderes  distortus, 
274. 

Mandibles,  use  of  the,  in  Ammo- 
phila,  273;  large,  of  Corydalis 
cornutus,  273;  large,  of  male  Lu- 
canus  elaphus,  273. 

Mandrill,  number  of  caudal  verte- 
brse  in  the,  56;  colors  of  the 
male,  534,  536,  545. 

Mantegazza,  Prof.,  on  last  molar 
teeth  of  man,  20;  bright  colors 
in  male  animals,  219;  on  the  or- 
naments of  savages,  568  et  seq. ; 
on  the  beardlessness  of  the  New 
Zealanders,  575;  on  the  exagger- 
ation of  natural .  cliaractfeTs  by 
man,  577. 


Mantell,  W.,  on  the  engrossment 
of  pretty  girls  by  the  New  Zea- 
land chiefs,  590, 

Mantis,  pugnacity  of  species  of, 
286. 

Maories,  mortality  of,  180;  infanti- 
cide and  proportion  of  sexes,  253; 
distaste  for  hairiness  amongst 
men,  575. 

Marcus  Aurelius,  on  the  origin  of 
the  moral  sense,  95;  on  the  influ- 
ence of  habitual  thoughts,  119. 

Mareca  penelope,  409. 

Marks,  retained  throughout 
groups  of  birds,  421. 

Marriage,  restraints  upon,  among 
savages,  44;  influence  of,  upon 
morals,  115;  influence  of,  on 
mortality,  136;  development  of, 
584. 

Marriages,  early,  134,  135;  com- 
munal, 582,  584. 

Marshall,  Dr.  W.,  protuberances 
on  birds'  heads,  235,  378;  on  the 
moulting  of  birds,  387;  advant- 
age to  older  birds  of  paradise, 
480. 

,  Col.,  interbreeding  amongst 

Todas,  184;  infanticide  and  pro- 
portion of  sexes  with  Todas,  252; 
choice  of  husbands  amongst  To- 
das, 588. 

,    Mr.,    on    the    brain    of    a 

Bushwoman,  163. 

Marsupials,  153;  development  of 
the  nictitating  membrane  in,  16; 
uterus  of,  37;  possession  of  nip- 
ples by,  158;  their  origin  from 
Monotremata,  160;  abdominal 
sacks  of,  204;  relative  size  of  the 
sexes  of,  511;  colors  of,  529. 

Marsupium,  rudimentary,  in  male 
marsupials,  157, 

Martin,  W.  C.  Lf.,  on  alarm  man- 
ifested by  an  orang  at  the  sight 
of  a  turtle,  71;  on  the  hair  in 
Hylobates,  148;  on  a  female 
American  deer,  510;  on  the  voice 
of  Hylobates  agihs,  523;  on  Sem- 
nopithecus  nemseus,  548. 

,  on  the  beards  of  the  in- 
habitants of  St.  Kilda,   555. 

Martins  deserting  their  young, 
104. 

,  C,  on  death  caused  by  in- 
flammation of  the  vermiform 
appendage,  20. 

Mastoid  processes  in  man  and 
apes,  51. 

Maudsley,  Dr.,  on  the  influence  of 
the  sense  of  smell  in  man,  IS: 
on  idiots  smelling  their  food,  35; 
on  Laura  Bridgman,  85;  on  the 
development  of  the  vocal  or- 
gans, 87;  moral  sense  failing  in 
incipient  madness,  120;  change 
of  mental  faculties  at  puberty 
in  man,  560. 

Mayers,  W.  F.,  on  the  domestica- 
tion o'f  the  gbldfish  in  China, 
339. 


INDEX. 


649 


Mayhew,  E.,  on  the  affection  be- 
tween individuals  of  different 
sexes  in  the  dog,  519, 

Maynard,  C.  J.,  on  the  sexes  of 
Chrysemys  picta,  346. 

Meckel,  on  correlated  variation  of 
the  muscles  of  the  arm  and  leg, 
42. 

Medicines,  effect  produced  by,  the 
same  in  man  and  in  monkeys,  7. 

Medusae,  bright  colors  of  some, 
257. 

Megalithic  structures,  prevalence 
of,  175. 

Megapicus  validus,  sexual  differ- 
ence of  color  in,  452. 

Megasoma,  large  size  of  males  of, 
276. 

Meigs,  Dr.  A.,  on  variation  in  the 
skulls  of  the  natives  of  Amer- 
ica, 25. 

Meinecke,  on  the  numerical  pro- 
portion of  the  sexes  in  butter- 
flies,  246. 

Melanesians,  decrease  of,  181. 

Meldola,  Mr.,  colors  and  mar- 
riage flight  of  Colias  and  Pieris, 
315. 

Meliphagidae,  Australian,  nidifi- 
cation  of,  448. 

Melita,  secondary  sexual  char- 
acters of,  265. 

Meloe,  difference  of  color  in  the 
sexes  of  a  species  of,  291. 

Memory,  manifestations  of,  in  an- 
imals, 72. 

Memnon,  young,  164. 

Mental  characters,  difference  of, 
in  different  races  of  men,  164. 

faculties,  diversity  of,  in  the 

same  race  of  men,  25;  inherit- 
ance of,  27;  variation  of,  in  the 
same  species,  27,  64;  similarity  of 
the,  in  different  races  of  man, 
174;  of  birds,  405. 

powers,  difference  of,  in  the 


two  sexes  in  man,  558. 
Menura  Alberti,  401;  song  of,  366. 

superba,    401;    long   tails   of 

both  sexes  of,  445. 

Merganser,  trachea  of  the    male, 
369. 

serrator,  male  plumage  of, 

388. 

Mergus  cucullatus,  speculum    of, 
231. 

merganser,  young  of,  462. 

Metallura,     splendid   tail-feathers 

of,  437. 
Methoca      ichneumonides,      large 

male  of,  277. 
Meves,  M.,  on  the  drumming    of 

the  snipe,  373. 
Mexicans,  civilization  of  the,  not 

foreign,  141. 
Meyer,   on   a  convoluted  body   at 

the   extremity  of   the    tail  in   a 

Macacus  and  a  cat,  22. 
,  Dr.  A.,  on  the  copulation  of 

Phryganidse   of  distinct  species, 

272. 


Meyer,  Prof.  L.,  on  development 
of  helix  of  ear,  15;  men's  ears 
more  variable  than  women's, 
219;  antennse  serving  as  ears, 
278. 

Migrations  of  man,  effects  of,  46. 

Migratory  instinct  of  birds,  101; 
vanquishing  the  maternal,  104, 
107. 

Mill,  J.  S.,  on  the  origin  of  the 
moral  sense,  95;  on  fhe  "greatest 
happiness  principle,"  116;  on  the 
difference  of  the  mental  powers 
in  the  sexes  of  man,  559. 

Millipedes,  270. 

Milne-Edwards,  H.,  on  the  use  of 
the  enlarged  chelae  of  the  male 
Gelasimus,  263. 

Milvago  leucurus,  sexes  and 
young  of,  473. 

Mimicry,   323. 

Mimus  polyglottus,  406. 

Mind,  difference  of,  in  man  and 
the  highest  animals,  122;  simi- 
larity of  the,  in  different  races, 
174. 

Minnow,  proportion  of  the  sexes 
in  the.  245. 

Mirror,   larks   attracted  by,   407. 

Mitchell,  Dr.,  interbreeding  in  thie 
Hebrides,  184. 

Mitford,  selection  of  children  in 
Sparta,  28. 

Mivart,  St.  George,  on  the  reduc- 
tion of  organs,  11;  on  the  ears 
of  the  lemuroidea,  15;  on  varia- 
bility of  the  muscles  in  lemuroi- 
dea, 40,  46;  on  the  caudal  verte- 
bras of  monkeys,  56;  on  the 
classiflcation  of  the  primates, 
149;  on  the  orang  and  on  man, 
149,  150;  on  differences  in  the 
lemuroidea,  150;  on  the  crest  of 
the  male  newt,  344. 

Mobius,  Prof.,  on  reasoning  pow- 
ers in  a  pike,  73. 

Mocking-thrush,  partial  migration 
of,  406;  young  of  the,  482. 

Modifications,   unserviceable,  59. 

Moggridge,  J.  T.,  on  habits  of 
spiders,   67;    on    habits    of    ants. 

Moles,  numerical  proportion  of 
the  sexes  in,  242;  battles  of  male, 

496. 

Mollienesia  petenensis,  sexual  dif- 
ference in,   334. 

Mollusca,  beautiful  colors  and 
shapes  of,  260;  absence  of  sec- 
ondary sexual  characters  in  the, 
259. 

Molluscoida,  155,  259. 

Monacanthus  scopas  and  M.  Per- 
onii,  328. 

Mongolians,  perfection  of  the 
senses  in,   33. 

Monkey,  protecting  his  keeper 
from  a  baboon,  100,  107;  bonnet-, 
146;  rhesus-,  sexual  difference  in 
color  of  the,  534,  546;  moustache-, 
colors  of  the.  532. 


650 


INDEX. 


Monkeys,  liability  of,  to  the  same 
diseases  as  man,  7;  male,  recog- 
nition of  women  by,  8;  diversity 
of  the  mental  faculties  in,  26; 
breaking  hard  fruits  with 
stones,  49;  hands  of  the,  48,  49; 
basal  caudal  vertebrae  of,  im- 
bedded in  the  body,  57;  revenge 
taken  by,  67;  maternal  affection 
in,  68;  variability  of  the  faculty 
of  attention  in,  71;  American, 
manifestation  of  reason  in,  75; 
using  stones  and  sticks,  79;  imi- 
tative faculties  of,  85;  signal- 
cries  of,  85;  mutual  kindnesses 
of,  98;  sentinels  posted  by,  97; 
human  characters  of,  146;  Amer- 
ican, direction  of  the  hair  on  the 
arms  of  some,  147;  gradation  of 
species  of,  171;  beards  of,  526; 
ornamental  characters  of,  543; 
analogy  of  sexual  differences  of, 
with  those  of  man,  553;  different 
degrees  of  difference  in  the 
sexes  of,  556;  expression  of  emo- 
tions by,  566;  generally  monog- 
amous habits  of,  585;  polyga- 
mous habits  of  some,  585;  naked 
surfaces  of,  595. 

Monogamy,   not  primitiye,   140. 

Monogenists,  171. 

Mononychus  pseudacori,  stridula- 
tion  of,  302. 

Monotremata,  153;  development  of 
the  nictitating  membrane  in,  17; 
lactiferous  glands  of,  158;  con- 
necting mammals  with  reptiles, 
160. 

Monstrosities,  analogous,  in  man 
and  lower  animals,  29;  caused 
by  arrest  of  development,  34; 
correlation  of,  42;  transmission 
of,  169. 

Montagu,  G.,  on  the  habits  of  the 
black  and  red  grouse,  215;  on 
the  pugnacity  of  the  ruff,  358; 
on  the  singing  of  birds,  364;  on 
the  double  moult  of  the  male 
pintail,  387. 

Monteiro,  Mr.,  on  Bucorax  abys- 
sinicus,  377. 

Montes  de  Oca,  M.,  on  the  pug- 
nacity of  male  Humming-birds, 
356. 

Monticola  cyanea,  450. 

Monuments,  as  traces  of  extinct 
tribes,  177. 

Moose,  battles  of,  497;  horns  of 
the,   an  incumbrance,   510. 

Moral  and  instinctive  impulses,  al- 
liance of,  108. 

faculties,  their  influence  on 

natural  selection  in  man,  124. 

rules,    distinction    between 


the  higher  and  lower,  118. 

sense,     so-called,     derived 


from    the    social    instincts,    117; 
origin  of  the,  120. 

tendencies,    inheritance    of, 


120. 


Morality,  supposed  to  be  founded 
in  selfishness,  116;  test  of,  the 
general  welfare  of  the  com- 
munity, 118;  gradual  rise  of,  121; 
influence  of  a  high  standard  of, 
129. 

Morgan,  L.  H.,  on  the  beaver,  65; 
on  the  reasoning  powers  of  the 
beaver,  73;  on  the  forcible  cap- 
ture of  wives,  140;  on  the  cas- 
toreum  of  the  beaver,  524;  mar- 
riage unknown  in  primeval 
times,  582;  on  polyandry,  587. 

Morley,  J.,  on  the  appreciation  of 
praise  and  fear  of  blame,  139. 

Morris,  F.  O.,  on  hawks  feeding 
an  orphan  nestling,  404. 

Morse,  Dr.,  colors  of  mollusca, 
260. 

Morselli,  E.,  division  of  the  malar 
bone,  37. 

Mortality,  comparative,  of  fe- 
males and  males,  212,  238. 

Morton,  on  the  number  of  species 
of  man,  170. 

Moschkau,  Dr.  A.,  on  a  speaking 
starling,  83. 

Moschus  moschiferus,  odoriferous 
organs  of,  525. 

Motacillse,   Indian,    young   of,    463. 

Moths,  310;  absence  of  mouth  in 
some  males,  204^^  apterous  fe- 
male, 204;  male,  prehensile  us© 
of  the  tarsi  by,  205;  male,  at- 
tracted by  females,  247;  colora- 
tion of,  312;  sexual  differences 
of  color  in,  812. 

Motmot,  inheritance  of  mutilation 
of  tail  feathers,  57,  597;  racket- 
shaped  feathers  in  the  tail  of  a, 
380. 

Moult,  double,  456;  double  annu- 
al,  in  birds,  385. 

Moulting  of  birds,  478. 

Moults,  partial,  386. 

Mouse,    song  of,   562. 

Moustache-monkey,  colors  of  the, 
532,   547. 

Moustaches,  in  monkeys,  146. 

Mud-turtle,  long  claws  of  the 
male,  346. 

Mulattoes,  persistent  fertility  of, 
167;  immunity  of,  from  yellow 
fever,   188. 

Mule,  sterility  and  strong  vitality 
of  the,   167. 

Mules,  rational,  76. 

Muller,  Ferd.,  on  the  Mexicans 
and  Peruvians,  141. 

,  Fritz,  on  astomatous  males 

of  Tanais,  204;  on  the  disappear- 
ance of  spots  and  stripes  in 
adult  mammals,  542;  on  the  pro- 
portions of  the  sexes  in  some 
Crustacea,  252;  on  secondary 
sexual  characters  in  various 
Crustaceans,  262  et  seq. ;  musical 
contest  between  male  Cicadas, 
280;  mode  of  holding  wings  in 
Castnia,    311;    on   birds   showing 


INDEX. 


651 


a  preference  for  certain  colors, 
313;  on  the  sexual  maturity  of 
young  amphipod  Crustacea,  479. 
Hermann,     emergence     of 


bees  from  pupa,  209;  pollen- 
gathering  of  bees,  223;  propor- 
tion of  sexes  in  bees,  250;  court- 
ing of  Eristalis,  278;  color  and 
sexual  selection  with  bees,  290. 
-,  J.,  on  the  nictitating  mem- 


brane and  semilunar  fold,  17. 

,  Max,  on  the  origin  of  lan- 
guage, 84;  language  implies 
power  of  general  conception,  86: 
struggle  for  life  among  the 
words,  &c.,  of  languages,  88. 

,   S.,  on  the  banteng,  532;  on 

the  colors  of  Semnopithecus 
chrysomelas,  533. 

Muntjac-deer,  weapons  of  the, 
509. 

Murie,  J.,  on  the  reduction  of 
organs,  11;  on  the  ears  of  the 
Lemuroidea,  15;  on  variability  of 
the  muscles  in  the  Lemuroidea, 
40,  46;  basal  caudal  vertebrae 
of  Macacus  brunneus  imbedded 
in  the  body,  57;  on  the  manner  of 
sitting  in  short-tailed  apes,  57; 
on  differences  in  the  Lemuroi- 
dea, 150;  on  the  throat-pouch  of 
the  male  bustard,  368;  on  the 
mane  of  Otaria  jubata,  516;  on 
the  sub-orbital  pits  of  Rumin- 
ants, 525;  on  the  colors  of  the 
sexes  in   Otaria  nigrescens,   530. 

Murray,  A.,  on  the  Pediculi  of 
different  races  of  men,   166. 

,    T.    A.,    on   the   fertility   of 

Australian  women  with  white 
men,  166. 

Mus  coninga,  78. 

minutus,     sexual   difference 

in  the  color  of,   529. 

Musca  vomitoria,  52. 
Muscicapa  grisola,   449. 

luctuosa,  449. 

ruticilla,  breeding  in  imma- 
ture plumage,  479. 

Muscle,   ischio-pubic,   40. 

Muscles,  rudimentary,  occurrence 
of,  in  man,  11;  variability  of  the, 
26;  effects  of  use  and  disuse 
upon,  31;  animal-like  abnormali- 
ties of,  in  man,  40;  correlated 
variation  of,  in  the  arm  and  leg, 
42;  variability  of,  in  the  hands 
and  feet,  46;  of  the  jaws,  influ- 
ence of,  on  the  physiognomy  of 
the  Apes,  52;  habitual  spasms 
of,  causing  modifications  of  the 
facial  bones,  53;  of  the  early 
progenitors  of  man,  156;  great- 
er variability  of  the,  in  men 
than  in  women,   219. 

Musculus  sternalis,  Prof.  Turner 
on  the,  13. 

Music,  174;  of  birds,  363;  discord- 
ant, love  of  savages  for,  375; 
reason  of  power  of  perception 
of  notes  in  animals,  563;  power 


of  distinguishing  notes,  564;  its 
connection  with  primeval 
speech,  567;  different  apprecia- 
tion of,  by  different  peoples,  564; 
origin  of,  564,  567;  effects  of,  566. 

Musical,  cadences,  perception  of, 
by  animals,  564;  powers  of  man, 
561  et  seq. 

Musk-deer,  canine  teeth  of  male, 
498,  509,  510;  male,  odoriferous 
organs  of  the,  525;  winter  change 
of  the,  537. 

Musk-duck,  Australian,  355;  large 
size  of  male,  358;  of  Guiana, 
pugnacity  of  the  male,  358. 

Musk-ox,  horns  of,  501. 

Musk-rat,  protective  resemblance 
of  the,  to  a  clod  of  earth,  537. 

Musophag^,  colors  and  nidifica- 
tion  of  the,  449;  both  sexes  of, 
equally  brilliant,  453. 

Mussels  opened  by  monkeys,  49. 

Mustela,  winter  change  of  two 
species  of,  537. 

Musters,  Capt.,  on  Rhea  Darwinii, 
473;  marriages  amongst  Pata- 
gonians,  592. 

Mutilations,  healing  of,  8;  inheri- 
tance of,  57. 

Mutillidse.  absence  of  ocelli  in  fe- 
male, 272. 

Mycetes  caraya,  polygamous,  213; 
vocal  organs  of,  523;  beard  of, 
527;  sexual  differences  of  color 
in,  532;   voice  of,  562. 

seniculus,  sexual  differences 

of  color  in,  532. 

Myriapoda,  270. 


N 


Nageli,  on  the  influence  of  nat- 
ural selection  on  plants,  58;  on, 
the  gradation  of  species  of 
plants,  171. 

Nails,  colored  yellow  or  purple  in 
part  of  Africa,  569. 

Narv/hal,   tusks  of  the,  498,   503. 

Nasal  cavities,  large  size  of,  in 
American  aborigines,   33. 

Nascent  organs,  11. 

Nathusius,  H.  von,  on  the  im- 
proved breeds  of  pigs,  173;  male 
domesticated  animals  more  vari- 
able than  females,  218;  horns 
of  castrated  sheep,  502;  on  the 
breeding  of  domestic  animals, 
590. 

Natural  selection,  its  effects  on 
the  ea.rly  progenitors  of  man, 
46;  influence  of  on  man,  58,  59; 
limitation  of  the  principle,  58; 
influence  of,  on  social  animals, 
60;  Mr.  Wallace  on  the  limita- 
tion of,  by  the  influence  of  the 
mental  faculties  in  man,  124;  in- 
fluence of,  in  the  progress  of  the 
United  States,  138;  in  relation  to 
sex,   255. 


652 


INDEX. 


Natural  and  sexual  selection  con- 
trasted, 221. 

Naulette,  jaw  from,  large  size  of 
the  canines  in,  39. 

Neanderthal  skull,  capacity  of 
the,   53. 

Neck,  proportion  of,  in  soldiers 
and  sailors,  31. 

Necrophorus,  stridulation  of,  299, 
301. 

Nectarinia,  young  of,  468. 

Nectarinise,  moulting  of  the,  392; 
nidification  of,  462. 

Negro,  resemblance  of  a,  to  Euro- 
peans, in  mental  characters,  174. 

Negro-women,  their  kindness  to 
Mungo   Park,    114. 

Negroes,  Caucasian  features  in, 
163;  character  of,  164;  lice  of,  166; 
fertility  of,  when  crossed  with 
other  races,  167;  blackness  of, 
166,  169;  variability  of,  170,  171; 
immunity  of,  from  yellow  fever, 
188;  difference  of,  from  Ameri- 
cans, 192;  disfigurements  of  the, 
536;  color  of  new-born  children 
of  552;  comparative  beardless- 
ness  of,  555;  readily  become 
musicians,  565;  appreciation  of 
beauty  of  their  women  by,  572; 
573;  idea  of  beauty  among,  576; 
compression  of  the  nose  by 
some,  577. 

Nemertians,  colors  of,  261. 

Neolithic  period,  141. 

Neomorpha,  sexual  difference  of 
the  beak  in,  356. 

Nephila,   size  of  male,  270. 

Nests,  made  by  fishes,  341;  decor- 
ation of,  by  Humming-birds, 
408. 

Neumeister,  on  a  change  of  color 
in  pigeons  after  several  moult- 
ings,  233. 

Neuration,  difference  of,  ^,n  the 
two  sexes  of  some  butterflies 
and  hymenoptera,  274. 

Neuroptera,  250,  287. 

Neurothemis,   dimorphism  in,  288. 

New  Zealand,  expectation  by  the 
natives  of,  of  their  extinction, 
186;  practice  of  tattooing  in,  571; 
aversion  of  natives  of,  to  hairs 
on  the  face,  575;  pretty  girls  en- 
grossed by  the  chiefs  in,  590. 

Newton,  A.,  on  the  throat-pouch 
of  the  male  bustard,  368;  on  the 
difference  between  the  females 
of  two  species  of  Oxynotus,  464; 
on  the  habits  of  the  Phalarope, 
dotterel,  and  godwit,  472. 

Newts,  344. 

Nicholson,  Dr.,  on  the  non-immu- 
nity of  dark  Europeans  from 
yellow  fever,  190. 

Nictitating  nmembrane,   16,   156. 

Nidification,  of  fishes,  340;  rela- 
tion of,  to  color,  447,  450;  of  Brit- 
ish birds,  448. 

Night-heron,   cries  of  the,   363. 


Nightingale,  arrival  of  the  male 
before  the  female,  208;  object  of 
the  song  of  the,  364. 

Nightingales,  new  mates  found  by 
403. 

Nightjar,  selection  of  a  mate  by 
the  female,  410;  Australian, 
sexes  of,  473;  coloration  of  the, 
486. 

Nightjars,  noise  made  by  some 
male,  with  their  wings,  371; 
elongated  feathers  in,  379,  397. 

Nilghau,  sexual  differences  of  col- 
or in  the,  530. 

Nilsson,  Prof.,  on  the  resem- 
blance of  stone  arrow-heads 
from  various  places,  174;  on  the 
development  of  the  horns  in  the 
reindeer,   229. 

Nipples,  absence  of,  in  Monotre- 
mata,  158. 

Nitsche,  Dr.,  ear  of  foetal  orang, 
16. 

Nitzsch,  C.  L.,  on  the  down  of 
birds,  385. 

Noctuse,  brightly-colored  be- 
neath, 312. 

Noctuidse,  coloration  of,  310. 

Nomadic  habits,  unfavorable  to 
human  progress,  130. 

Nordmann,  A.,  on  Tetrao  urogal- 
loides,  400. 

Norfolk  island,  half-breeds  on,  186. 

Norway,  numerical  proportion  of 
male  and  female  births  in,  237. 

Nose,  resemblance  of,  in  man  and 
the  apes,  149;  piercing  and  orna- 
mentation of  the,  570;  very  flat, 
not  admired  in  negroes,  577;  flat- 
tening of  the,  577. 

Nott  and  Gliddon,  on  the  features 
of  Rameses  II.,  164;  on  the  fea- 
tures of  Amunoph  III.,  164;  on 
skulls  from  Brazilian  caves,  165: 
on  the  immunity  of  negroes  and 
mulattoes  from  yellow  fever,  188; 
on  the  deformation  of  the  skull 
among  American  tribes,  577. 

Novara,  voyage  of  the,  suicide  in 
New  Zealand,  113. 

Nudibranch  Mollusca,  bright  col- 
ors of,  260. 

Ntimerals,   Roman,  140. 

Nunemaya,  natives  of,  bearded, 
555. 

Nut-hatch  of  Japan,  Intelligence 
of,  405. 


O 


Obedience,  value  of,  127. 
Observation,  powers  of,  possessed 

by  birds,  406. 
Occupations,    sometimes    a    cause 

of  diminished  stature,  30;   effect 

of,  upon  the  proportions  of  the 

body,  30. 
Ocelli,  absence  of,  in  female  Mut= 

illidae,  272. 


INDEX. 


653 


Ocelli  of  birds,  formation  and  va- 
riability of  the,  422. 

Ocelot,  sexual  differences  in  the 
coloring  of  the,  529. 

Ocyphaps  lophotes,  396. 

Odonata,  250. 

Odonestis  potatoria,  sexual  differ- 
ence of  color  in,  312. 

Odor,  correlation  of,  with  color  of 
skin,  192;  emitted  by  snakes  in 
the  breeding-season,  348;  of 
mammals,  524. 

Oecanthus  nivalis,  difference  of 
color  in  the  sexes  of,  287. 

pellucidus,  287. 

Ogle,  Dr.  W.,  relation  between 
color  and  power  of  smell,  18. 

Oidemia,   486. 

Oliver,  on  sounds  produced  by 
Pimelia  striata,  303. 

Omaloplia  brunnea,  stridulation 
of,  300. 

Onitis,  furcifer,  processes  of  ante- 
rior femora  of  the  male,  and  on 
the  head  and  thorax  of  the  fe- 
male,  295, 

Onthophagus,  293. 

rangifer,   sexual  differences 

of,  294;  variation  in  the  horns  of 
the  male,    295. 

Ophidia,  sexual  differences  of,  357. 

Ophidium,  343. 

Opossum,  wide  range  of,  in  Amer- 
ica, 165. 

Optic  nerve,  atrophy  of  the, 
caused  by  destruction  of  the  eye, 
31. 

Orang-Outan,  556;  Bischoff  on  the 
agreement  of  the  brain  of  the, 
with  that  of  man,  6;  adult  age 
of  the,  8;  ears  of  the,  14;  ver- 
miform appendage  of,  20;  hands 
of  the,  48;  absence  of  mastoid 
processes  in  the,  51;  platforms 
built  by  the,  64;  alarmed  at  the 
sight  of  a  turtle,  71;  using  a 
stick  as  a  lever,  79;  using  mis- 
siles, 79;  using  the  leaves  of  the 
Pandanus  as  a  night  covering, 
80;  direction  of  the  hair  on  the 
arms  of  the,  147;  its  aberrant 
characters,  149;  supposed  evolu- 
tion of  the,  173;  voice  of  the,  522; 
monogamous  habits  of  the,  585; 
male,  beard  of  the,  526. 

Oranges,    treatment  of,    by  mon- 

Orange-tip  butterfly,   305,   308,   309. 
Orchestia   Darwinii,     dimorphism 
of  males  of,  265. 

Tucuratinga,    limbs   of,   263, 

268. 

Ordeal,  trial  by,  93. 

Oreas  canna,  colors  of,  531. 

• Derbianus,     colors   of,    531, 

538. 

Organs,  prehensile,  205;  utilized 
for  new  purposes,  565. 

Organic  scale,  von  Baer's  defini- 
tion of  progress  in,  159. 


Orioles,  nidification  of,  447. 
Oriolus,    species    of,    breeding    in 
immature  plumage,  479. 

melanocephalus,    coloration 

of  the  sexes  in,  454. 

Ornaments,  prevalence  of  similar, 

175;  of  male  birds,  363;  fondness 

of  savages  for,  568. 
Ornamental      characters,       equal 

transmission  of,   to  both   sexes, 

in  mammals,     536;   of  monkeys, 

543. 
Ornithoptera  craesus,  246. 
Ornithorhynchus,     152;       reptilian 

tendency   of,   154;      spur   of   the 

male,  498. 
Orocetes  erythrogastra,  young  of, 

482. 

Orrony,  Grotto  of,  21. 

Orsodacna  atra,  difference  of  color 
in  the  sexes  of,  291. 

Orthoptera,  280;  metamorphosis 
of,  231;  stridulating  apparatus 
of,  280,  284;  colors  of,  286;  rudi- 
mentary stridulating  organs  in 
female,  284;  stridulation  of  the, 
and  Homoptera,   discussed,  286. 

Ortygornis  gularis,  pugnacity  of 
the  male,   359. 

Oryctes,  stridulation  of,  300;  sex- 
ual differences  in  the  stridulant 
organs  of,  302. 

Oryx  leucoryx,  use  of  the  horns 
of,  505,  513. 

Osphranter  rufus,  sexual  differ- 
ence in  the  color  of,  529. 

Ostrich,  African,  sexes  and  incu- 
bation of  the,  473. 

Ostriches,  stripes  of  young,  459. 

Otaria  jubata,  mane  of  the  male, 
516. 

nigrescens,  difference  in  the 

coloration  of  the  sexes  of,  530. 

Otis  bengalensis,  love-antics  of 
the  male,  376. 

tarda,  throat-pouch  of    the 

male,  368;  polygamous,  215. 

Ouzel,  ring-,  colors  and  nidifica- 
tion of  the,  449. 

,  water-,  colors  and  nidifica- 
tion of  the,  449. 

Ovibos  moschatus,  horns  of,  501. 

Ovipositor  of  insects,  204. 

Ovis  cycloceros,  mode  of  fighting 
of,   504,  509. 

Ovule  of  man,  7. 

Owen,  Prof.,  on  the  Corpora 
Wolfliana,  11;  on  the  great  toe  in 
man,  11;  on  the  nictitating 
membrane  and  semilunar  fold, 
17;  on  the  development  of  the 
posterior  molars  in  different 
races  of  man,  20;  on  the  length 
of  the  coecum  in  the  Koala,  20; 
of  the  coccygeal  vertebrae,  22; 
on  rudimentary  structures  be- 
longing to  the  reproductive  sys- 
tem, 23;  on  abnormal  conditions 
of  the  human  uterus,  37;  on  the 
number  of  digits  in  the  Ichthy- 


654 


INDEX. 


©pterygia,  35;on  the  canine  teeth 
in  man,  39;  on  the  walking  of 
the  chimpanzee  and  orang,  48; 
on  the  mastoid  processes  in  the 
higher  apes,  51;  on  the  hairmess 
of  elephants  in  elevated  dis- 
tricts, 55;  on  the  caudal  verte- 
brae of  monkeys,  56;  classifica- 
tion of  mammalia,  144;  on  the 
hair  in  monkeys,  148;  on  the  pis- 
cine affinities  of  the  Ichthyos- 
aurians,  154;  on  polygamy  and 
monogamy  among  the  antelopes, 
213;  on  the  horns  of  Antilocapra 
americana,  229;  on  the  musky 
odor  of  crocodiles  during  the 
breeding  season,  347;  on  the 
scent-glands  of  snakes,  348;  on 
the  Dugong,  Cachalot  and  Or- 
nithorhynchus,  498;  on  the  ant- 
lers of  the  red  deer,  506;  on  the 
dentition  of  the  Camelidas,  509; 
on  the  horns  of  the  Irish  elk, 
511;  on  the  voice  in  the  giraffe, 
porcupine,  and  stag,  521;  on  the 
laryngeal  sac  of  the  gorilla  and 
orang,  522;  on  the  odoriferous 
glands  of  mammals,  529;  on  the 
effects  of  emasculation  on  the 
vocal  organs  of  men,  566;  on  the 
voice  of  Hylobates  agilis,  561;  on 
American  monogamous  mon- 
keys, 585. 
Owls,  white,  new  mates  found  by, 

403.  .    .^      X 

Oxynotus,    difference    of    the    fe- 
males of  two  species  of,  464. 


Pachydermata,  213. 

Pachytylus  migratorius,  280. 

Paget,  on  the  abnormal  develop- 
ment of  hairs  in  man,  18;  on  the 
thickness  of  the  skin  on  the 
soles  of  the  feet  of  infants,  32. 

Painting,  pleasure  of  savages  in, 
174. 

Palaemon,  chelae  of  a  species  or, 
263. 

Palaeornis,  sexual  differences  of 
color  in,   489. 

javanicus,  color  of  beak  or, 


455. 


rosa,  young  of,  461. 


Palamedea  cornuta,  spurs  on  the 
wings,   360. 

Paleolithic  period,  141. 

Palestine,  habits  of  the  chaffinch 
in,  244. 

Pallas,  on  the  perfection  of  the 
senses  in  the  Mongolians,  33;  on 
the  want  of  connection  between 
chmate  and  the  color  of  the 
skin,  187;  on  the  polygamous 
habits  of  Antilope  Saiga,  213;  on 
thp  lig-hter  color  of  horses  and 
cattle  in  winter  in  Siberia,  224; 
on  the  tusks  of  the  musk-deer, 


509;  on  the  odoriferous  glands  of 
mammals,  525;  on  the  odorifer- 
ous glands  of  the  musk-deer, 
525;  on  winter  changes  of  color 
in  mammals,  537;  on  the  ideal  of 
female  beauty  in  North  China, 
572. 

Palmaris  accessorius,  muscle  va- 
riations of  the,  26. 

Pampas,   horses  of  the,  177. 

Pangenesis,  hypothesis  of,  223,  227. 

Panniculus  carnosus,   13. 

Pansch,  on  the  brain  of  a  foetal 
Cebus  apella,  201. 

Papilio,  proportion  of  the  sexes 
in  North  American  species  of, 
246;  sexual  differences  of  color- 
ing in  species  of,  306;  coloration 
of  the  wings  in  species  of,  309. 

ascanius,   307. 

Sesostris      and      Childrense, 

variability  of,  316. 

Turnus,  246. 

Papilionidse,     variability    in     the, 

316. 
Papuans,    line    of    separation   be- 
tween the,  and  the  Malays,  165; 
beards  of  the,  555;  hair  of,  569. 

and     Malays,     contrast    in 

characters  of,  164. 

Paradise,  Birds  of,  400,  456;  sup- 
posed by  Lesson  to  be  polygam- 
ous, 214;  rattling  of  their  quills 
by,  370;  racket-shaped  feathers 
in,  379;  sexual  differences  in 
color  of,  380;  decomposed  feath- 
ers in,  380,  397;  display  of  plu- 
mage by  the  male,  391. 

Paradisea  apoda,  barbless  feath- 
ers in  the  tail  of,  380;  plumage 
of,  380;  and  P.  papuana,  380;  di- 
vergence of  the  females  of,  464; 
increase  of  beauty  with  age, 
480. 

Paraguay,  Indians  of,  eradication 
of  eyebrows  and  eyelashes  by, 
576. 

Parakeet,  Australian,  variation  in 
the  color  of  the  thighs  of  a 
male,    418. 

Parallelism  of  development  of 
species  and  languages,  87. 

Parasites  on  man  and  animals,  7, 
8;  as  evidence  of  specific  identity 
or  distinctness,  165;  immunity 
from,  correlated  with  color,  188. 

Parental  feeling  in  earwigs,  star- 
fishes, and  spiders,  102;  affection, 
partly  a  result  of  natural  selec- 
tion, 102. 

Parents,  age  of,  influence  upon 
sex  of  offspring,  239. 

Parinae,  sexual  difference  of  color 
in,  451. 

Park,        Mungo,  negro-women 

teaching  their  children  to  love 
the  truth,  114;  his  treatment  by 
the  negro-women,  114:  on  negro 
opinions  of  the  appearance  of 
white  men,  573. 


INDEX. 


655 


Parker,  Mr.,  no  bir^  or  reptile  in 
line  of  mammalian  descent,  153. 

Parrot,  racket-shaped  feathers  in 
the  tail  of  a,  37^;  instance  of 
benevolence  in  a,  406. 

Parrots,  change  of  color  in,  58; 
imitative  faculties  of,  71;  living 
in  triplets,  404;  affection  of,  405; 
colors  and  nidification  of  the, 
449,  451,  452;  immature  plumage 
of  the,  461;  colors  of,  485;  sex- 
ual differences  of  color  in,  489; 
musical  powers  of,  565. 

Parthenogenesis  in  the  Tenthre- 
dinae,  251;  in  Cynipidae,  250;  in 
Crustacea,  251. 

Partridge,  monogamous,  248;  pro- 
portion of  the  sexes  in  the,  243; 
female,  465. 

" dances,"  375,  400. 

Partridges,  living  in  triplets,  404; 
spring  coveys  of  male,  404;  dis- 
tinguishing persons,  406. 

Parus  casruleus,  451. 

Passer,  sexes  and  young  of,  477. 

brachydactylus,  477. 

domesticus,  449,  477. 

montanus,  449,  477. 

Patagonians,       self-sacrifice     by, 

107;   marriages  of,  592. 
Patterson,  Mr.,  on  the  Agrionidse, 

288. 
Patteson,     Bishop,      decrease     of 

Melanesians,  181. 
Paulistas  of  Brazil,  169. 
Pavo  cristatus,  230,  424. 

muticus,  230,  424;  possession 

of  spurs  by  the  female,  360,  444. 

nigripennis,  413. 


Payaguas  Indians,  thin  legs  and 
thick  arms  of  the,  31. 

Payan,  Mr.,  on  the  proportion  of 
the  sexes  in  sheep,  219. 

Peacock,  polygamous,  214;  sexual 
characters  of,  230;  pugnacity  of 
the,  360;  rattling  of  the  quills 
by,  371;  elongated  tail-coverts  of 
the,  379,  397;  love  of  display  of 
the,  389,  425;  ocellated  spots  of 
the,  424;  inconvenience  of  long 
tail  of  the,  to  the  female,  438, 
445,  446;  continued  increase  of 
beauty  of  the,  480. 

butterfly,  309. 

Peafowl,  preference  of  females 
for  a  particular  male,  413;  first 
advances  made  by  the  female, 
415. 

Pediculi  of  domestic  animals  and 
man,  166. 

Pedigree  of  man,  161. 

Pedionomus  torquatus,  sexes  of, 
469. 

Peel,  J.,  on  horned  sheep,  501. 

Peewit,  wing-tubercles  of  the 
male,  361. 

Pelagic  animals,  transparency  of, 
259. 

Pelecanus  erythrorhynchus, horny 
crest  on  the  beak  of  the  male, 
during  the  breeding  season,  384. 


Pelecanus  onocrotalus,  spring 
plumage  of,  388. 

Pelele,  an  African  ornament,  570. 

Pelican,  blind,  fed  by  his  compan- 
ions, 99;  young,  guided  by  old 
birds,  99;  pugnacity  of  the  male, 
358. 

Pelicans,  fishing  in  concert,  98. 

Pelobius  Hermanni,  stridulation 
of,   300,   301. 

Pelvis,  alteration  of,  to  suit  the 
erect  attitude  of  man,  51;  differ- 
ences of  the,  in  the  sexes  in 
man,  552. 

Penelope  nigra,  sound  produced 
by  the  male,  373. 

Pennant,  on  the  battles  of  seals, 
496;  on  the  bladder-nose  seal, 
523. 

Penthe,  antennal  cushions  of    the 

male,  273. 
Perch,  brightness  of  male,  during 

breeding  season,  336. 
Peregrine  falcon,  new  mate  found 

by,  402. 
Period  of  variability,  relation  of, 

to  sexual  selection,  235. 
Periodicity,   vital.      Dr.   Laycock, 

on,   8. 

Periods,  lunar,  followed  by  func- 
tions in  man  and  animals,  8,  160. 

of  life,  inheritance  at  corre- 
sponding, 223,  227. 

Perisoreus  canadensis,  young  of, 
476. 

Peritrichia,  difference  of  color  in 
the  sexes  of  a  species  of,  292. 

Periwinkle,  259. 

Pernis  cristata,  418. 

Perrier,  M.,  on  sexual  selection, 
205;  on  bees,  290. 

Perseverance,  a  characteristic  of 
man,  558. 

Persians,  said  to  be  improved  by 
intermixture  with  Georgians 
and  Circassians,  581. 

Personnat,  M.,  on  Bombyx  Yam- 
amai,  246. 

Peruvians,  civilization  of  the,  not 
foreign,  141. 

Petrels,  colors  of,  489. 

Petrocincla  cyanea,  young  of,  482. 

Petrocossyphus,  455. 

Petronia,  477. 

Pfeiffer,  Ida,  on  Javan  ideas  of 
beauty,  574. 

Phacochcerus  sethiopicus,  tusks 
and  pads  of,  515. 

Phalanger,  Vulpine,  black  varie- 
ties of  the,  535. 

Phalaropus  fulicarius,  471. 

hyperboreus,  471. 

Phan^us,  295. 

—. carnifex,     variation   of   the 

horns  of  the  male,  294. 

faunus,     sexual  differences 


of,  293. 

lancifer,  293. 


Phaseolarctus  cinereus,  taste  for 
rum  and  tobacco,  7. 


656 


INDEX. 


Phasgonura  viridJ.ssima,  stridula- 
tion  of,  2S2,   2S3. 

i'hasianus  Scemmerringii,  440. 

versicolor,  391. 

. Wallichii,  395,  466. 

Pheasant,  polygamous,  215;  and 
black  grouse,  hybrids  of,  409; 
production  of  hybrids  with  the 
common  fowl,  414;  immature 
plumage  of  the,  461. 

Pheasant,  Amherst,  display  of, 
390. 

,   Argus,    379,   456;   display  of 

plumage  by  the  male,  392;  ocel- 
lated  spots  of  the,  422,  428;  grad- 
ation of  characters  in  the,  428. 

,  Blood-,  360. 

,  Cheer,  395,  467. 

Eared,   230,   395,   467;   length 


of    the    tail    in    the,    446;    sexes 
alike  in  the,  454. 

-,  Golden,  display  of  plumage 


by  the  male,  390;  age  of  mature 
plumage  in  the,  478;  sex  of 
young,  ascertained  by  pulling 
out  headfeathers,  478. 

Kalij,     drumming    of     the 


male,  371. 

Reeve's,   length  of  the  tail 


in,  446. 

-,  Silver,  triumphant  male,  de- 


posed on  account  of  spoiled  plu- 
mage, 413;  sexual  coloration  of 
the,  488. 

,  Soemmerring's,  439,  446. 

,   Tragopan,   377;    display     of 

plumage  by  the  male,  392;  mark- 
ings  of   the   sexes  of  the,   422. 

Pheasants,  period  of  acquisition  of 
male  characters  in  the  family  of 
the,  230;  proportion  of  sexes  in 
chicks  of,  242;  length  of  the  tail 
in,  439,   445,  446. 

Philters,   worn  by  women,  572. 

Phoca  grcEnlandica,  sexual  differ- 
ence in  the  coloration  of,  530. 

Phcenicura  ruticilla,  403. 

Phosphorescence  of  insects,  275. 

Phryganidse,  copulation  of  dis- 
tinct species  of,  272. 

Phryniscus  nigricans,  345. 

Physical  inferiority,  supposed,  of 
man,   60,    61. 

Pickering,  on  the  number  of  spe- 
cies of  man,  170. 

Picton,  J.  A.,  on  the  soul  of  man, 
607. 

Picus  auratus,  358. 

major,  396. 

Pieris,  309,  315. 

Pigeon,  female,  deserting  a 
weakened  mate,  210;  carrier,  late 
development  of  the  wattle  in, 
232;  pouter,  late  development  of 
the  crop  in,  232;  domestic,  breeds 
and  sub-breeds  of.  454. 

Pigeons,  nestling,  fed  by  the  se- 
cretion of  the  crop  of  both  pa- 
rents, 158;  changes  of  plumage 
in,  224;  transmission  of  sexual 
peculiarities     in,     225;     Belgian, 


with  black-streaked  males,  226, 
233,  440;  changing  color  after  sev- 
eral moultings,  233;  numerical 
proportion  of  the  sexes  in,  242; 
cooing  of,  370;  variations  in 
plumage  of,  380;  display  of  plu- 
mage by  male,  396;  local  memory 
of,  405;  antipathy  of  female,  to 
certain  males,  412;  pairing  of, 
412;  profligate  male  and  female, 
412;  wing-bars  and  tail-feathers 
of,  421;  supposititious  breed  of, 
439;  pouter  and  carrier,  peculiar- 
ities of,  predominant  in  males, 
441;  nidification  of,  447;  Austra- 
lian, 452;  immature  plumage  of 
the,  461. 

Pigs,  origin  of  the  improved 
breeds  of,  173;  numerical  propor- 
tion of  the  sexes  in,  241;  stripes 
of  young,  458,  541;  tusks  of  mi- 
ocene,  516;  sexual  preference 
shown  by,  520. 

Pike,  American,  brilliant  colors 
of  the  male,  during  the  breeding 
season,    337. 

,    reasoning     powers    of,    73; 

male,  devoured  by  females,  244. 

Pike,  L.  O.,  on  the  psychical  ele- 
ments of  religion,  93. 

Pimelia  striata,  sounds  produced 
by  the  female,  303. 

Pinel,  hairiness  in  idiots,  35. 

Pintail,  drake,  plumage  of,  388; 
pairing  with  a  wild  duck,  409. 

Duck,  pairing  with  a  wig- 
eon,    409. 

Pipe-fish,  filamentous,  340;  mar- 
supial receptacles  of  the  male, 
342. 

Pipits,    moulting   of   the,    386. 

Pipra,  modified  secondary  wing- 
feathers  of  male,  374. 

deliciosa,  374,  375. 

Pirates  stridulus,  stridulation  of, 
279. 

Pitcairn  island,  half-breeds  on, 
186. 

Pithecia  leucocephala,  sexual  dif- 
ferences of  color  in,  532. 

Satanas,  beard  of,  527,  528; 

resemblance  of,  to  a  negro,  598. 

Pits,  suborbital,  of  Ruminants, 
525. 

Pittidse,  nidification  of,  447. 

Placentata,  153. 

Plagiostomous  fishes,  327. 

Plain-wanderer,  Australian,  469. 

Planariae,  bright  colors  of  some, 
257. 

Plantain-eaters,  colors  and  nidi- 
fication of  the,  449;  both  sexes 
of,  equally  brilliant,  453. 

Plants,  cultivated,  more  fertile 
than  wild,  43;  Nageli,  on  natural 
selection  in,  58;  male  flowers  of, 
mature  before  the  female,  208; 
phenomena  of  fertilization  in, 
218. 

Platalea,  370;  change  of  plumage 
in,  455. 


INDEX. 


657 


Platyblemnus,  287. 

Platycercus,  young  of,  475. 

Platyphyllum  concavum,   280,   284. 

Platyrrhine  monkeys,  149. 

Platysma  rayoid.es,  13. 

Plecostomus,  head-tentacles  of 
the  males  of  a  species  of,  334. 

" barbatus,  peculiar  beard  of 

the  male,  334. 

Plectropterus  gambensis,  spurred 
wings  of,  360. 

Ploceus,   366,  371,  400. 

Plovers,  wing-spurs  of,  360;  double 
moult  in,  386. 

Plumage,  changes  of,  inheritance 
of,  by  fowls,  224;  tendency  to  an- 
alogous variation  in,  380;  display 
of,  by  male  birds,  388,  397; 
changes  of,  in  relation  to  sea- 
son, 456;  immature,  of  birds, 
459,  460;  color  of,  in  relation  to 
protection,  484. 

Plumes  on  the  head  in  birds, 
difference  of,  in  the  sexes,  445. 

Pneumora,    structure    of,    284. 

Podica,  sexual  difference  in  the 
color  of  the  irides  of,  419. 

Poeppig,  on  the  contact  of  civil- 
ized and  savage  races,  178. 

Poison,  avoidance  of,  by  animals, 
77. 

Poisonous  fruits  and  herbs  avoid- 
ed by  animals,  64. 

Poisons,  immunity  from,  corre- 
lated with  color-,  188. 

Polish  fowls,  origin  of  the  crest 
in,  226. 

Pollen  and  van  Dam,  on  the  colors 
of.  Lemur  macaco,  532. 

Polyandry,  587;  in  certain  Cypri- 
nidas,  245;  among  the  Elateridse, 
249. 

Polydactylism  in  man,  35. 

Polygamy,  influence  of,  upon  sex- 
ual selection,  212;  superinduced 
by  domestication,  215;  supposed 
increase  of  female  births  by,  239; 
in  the  stickleback,  328. 

Polygenists,   171. 

Polynesia,  prevalence  of  infanti- 
cide in,  586. 

Polynesians,  wide  geographical 
range  of,  28;  difference  of  stat- 
ure among  the,  30;  crosses  of, 
169;  variability  of,  170;  hetero- 
geneity of  the,  187;  aversion  of, 
to  hairs  on  the  face,  575. 

Polyplectron,  number  of  spurs  in, 
360;  display  of  plumage  by  the 
male,  392;  gradation  of  char- 
acters in,  425;  female  of,  465. 

chinquis,  392,  426. 

Hardwickii,  426. 

malaccense,  426,  427. 

Napoleonis,  426,  427. 

Polyzoa,  259. 
Pontoporeia  afflnis,  262. 
Porcupine,   mute,      except  in   the 

rutting  season,  521. 


Pores,  excretory,  numerical  rela- 
tion of,  to  the  hairs  in  sheep, 
198. 

Porpitee,  bright  colors  of  some, 
257. 

Portax  picta,  dorsal  crest  and 
throat- tuft  of,  526;  sexual  differ- 
ences of  color  in,  530,  538. 

Portunus  puber,  pugnacity  of,  265. 

Potamochoerus  penicillatus,  tusks 
and  facial  knobs  of  the,  516. 

Pouchet,  G.,  the  relation  of  in- 
stinct to  intelligence,  65;  on  the 
instincts  of  ants,  143;  on  the 
caves  of  Abou-Simbel,  164;  on 
the  immunity  of  negroes  from 
yellow  fever,  189;  change  of 
color  in  fishes,  340. 

Pouter  pigeon,  late  development 
of  the  large  crop  in,  232. 

Powell,  Dr.,  on  stridulation,  279. 

Power,  Dr.,  on  the  different  colors 
of  the  sexes  in  a  species  of 
Squilla,   267. 

Powys,  Mr.,  on  the  habits  of  the 
chaffinch  in  Corfu,  244. 

Pre-eminence  of  man,  46. 

Preference  for  males  by  female 
birds,  408,  415;  shown  by  mam- 
mals, in  pairing,  517. 

Prehensile  organs,  205. 

Presbytis  entellus,  fighting  of  the 
male,  557. 

Preyer,  Dr.,  on  function  of  shell 
of  ear,  14;  on  supernumerary 
mammae   in   women,    85. 

Prichard,  on  the  difference  of 
stature  among  the  Polynesians, 
30;  on  the  connection  between 
the  breadth  of  the  skull  in  the 
Mongolians  and  the  perfection 
of  their  senses,  33;  on  the  capac- 
ity of  British  skulls  of  different 
ages,  53;  on  the  flattened  heads 
of  the  Colombian  savages,  569; 
on  Siamese  notions  of  beauty, 
572;  on  the  beardlessness  of  the 
Siamese,  585;  on  the  deforma- 
tion of  the  head  among  Ameri- 
can tribes  and  the  natives  of 
Arakhan,   577. 

Primary  sexual  organs,  203. 

Primates,  145,  201;  sexual  differ- 
ences of  color  in,  532. 

Primogeniture,  evils  of,  132. 

Prionidse,  difference  of  the  sexes 
in  color,   291. 

Proctotretus  multimaculatus,   358. 

■  tenuis,   sexual  difference  in 

the  color  of,  354. 

Profligacy,  134. 

Progenitors,  early,  of  man,  156. 

Progress,  not  the  normal  rule  in 
human  society,  130;  elements  of, 
137. 

Prong-horn  antelope,  horns  of, 
229. 

Proportions,  difference  of,  in  dis- 
tinct races,  163. 


658 


INDEX. 


Protective  coloring  in  butterflies, 

308;  in  lizards,  354;   in.  birds,  467, 

484;  in  mammals,  537. 
nature  of  the  dull  coloring 

of  female   Lepidoptera,   317,   318, 

321. 

resemblances  in  fishes,   340. 


Protozoa,  absence  of  secondary- 
sexual  characters  in,  257. 

Pruner-Bey,  on  the  occurrence  of 
the  supra-condyloid  foramen  in 
the  humerus  of  man,  22;  on  the 
color  of  negro  infants,  553. 

Prussia,  numerical  proportion  of 
male  and  female  births   in,  237. 

Psocus,  proportions  of  the  sexes 
in,  251. 

Ptarmigan,  monogamous,  215; 
summer  and  winter  plumage  of 
the,  385,  386;  nuptial  assemblages 
of,  400;  triple  moult  of  the,  456; 
protective  coloration  of,  468. 

Puff -birds,  colors  and  nidification 
of  the,  449. 

Pugnacity  of  fine-plumaged  male 
birds,  395. 

Pumas,  stripes  of  young,  458. 

Puppies  learning  from  cats  to 
clean  their  faces,  71. 

P5'cnonotus,  hserhorrhous,  pug- 
nacity of  the  male,  357;  display 
of  under  tail  coverts  by  the 
male,  396. 

Pyranga  aestiva,  male  aiding  in 
incubation,  447. 

Pyrodes,  difference  of  the  sexes 
in  color,  291. 


Q 

Quadrumana,  hands  of,  48;  differ- 
ences between  man  and  the,  145; 
sexual  differences  of  color  in, 
532;  ornamental  characters  of, 
543;  analogy  of  sexual  differ- 
ences of,  with  those  of  man, 
553;  fighting  of  males  for  the 
females,  557;  monogamous  habits 
of,   585;    beards  of  the,    596. 

Quain,  R.,  on  the  variation  of  the 
muscles  in  man,  26. 

Quatrefages,  A.  de,  on  the  oc- 
currence of  a  rudimentary  tail 
in  man,  22;  on  variability,  29; 
on  the  moral  sense  as  a  distinc- 
tion between  man  and  animals, 
94;  civilized  men  stronger  than 
savages,  132;  on  the  fertility  of 
Australian  women  with  whitie 
men,  166;  on  the  Paulistas  of 
Brazil,  169;  on  the  evolution  of 
the  breeds  of  cattle,  173;  on  the 
Jews,  188;  on  the  liability  of 
negroes  to  tropical  fevers  after 
residence  in  a  cold  climate,  189; 
on  the  difference  between  fleld- 
and  house-slaves,  191;  on  the  in- 
fluence of  climate  on  color,  191; 
colors   of  annelids,   261;   on   the 


Ainos,  555;  on  the  women  of  San 
Giuliano,  581. 

Querquedula  acuta,  409. 

Quetelet,  proportion  of  sexes  in 
man,  237;  relative  size  in  man 
and  woman,  238. 

Quichua  Indians,  33;  local  varia- 
tion of  color  in  the,  191;  no  gray 
hair  among  the,  554;  hairlessness 
of  the,  556;  long  hair  of  the,  574. 

Quiscalus  major,  220;  proportions 
of  the  sexes  of,  in  Florida  and 
Honduras,  244. 


R 

Rabbit,  white  tail  of  the,  537. 

Rabbits,  domestic,  elongation  of 
the  skull  in,  54;  modification  of 
the  skull  in,  by  the  lopping  of 
the  ear,  54;  danger-signals  of, 
97;  numerical  proportion  of  the 
sexes  in,  241. 

Races,  distinctive  characters  of, 
163,  164;  or  species  of  man,  164; 
crossed,  fertility  or  sterility  of, 
166;  of  man,  variability  of  the, 
170;  of  man,  resemblance  of,  in 
mental  characters,  174;  forma- 
tion of,  176;  of  man,  extinction 
of,  177;  effects  of  the  crossing 
of,  187;  of  man,  formation  of  the, 
187;  of  man,  children  of  the,  552; 
beardless,  aversion  of,  to  hairs 
on  the  face,  574. 

Raffles,  Sir  S.,  on  the  banteng, 
532. 

Rafts,  use  of,  47,  175. 

Rage,  manifested  by  animals,  67. 

Raia  batis,  teeth  of,  331. 

clavata,    female    spined    on 

the  back,   327;  sexual  difference 

in  the  teeth  of,  331. 

maculata,  teeth  of,  331. 


Rails,  spur-winged,  360. 

Ram,  mode  of  fighting  of  the,  504; 
African,  mane  of  an,  528;  fat- 
tailed,  528. 

Rameses  II.,   features  of,  164. 

Ramsay,  Mr.,  on  the  Australian 
musk-duck,  355;  on  the  regent 
bird,  408;  on  the  incubation  of 
Menura  superba,  445. 

Rana  esculenta,  vocal  sacs  of,  346. 

Rat,  common,  general  dispersion 
of,  a  consequence  of  superior 
cunning,  78;  supplantation  of  the 
native,  in  New  Zealand,  by  the 
European  rat,  186;  common,  said 
to  be  polygamous,  214;  numeri- 
cal proportion  of  the  sexes  in, 
241. 

Rats,  enticed  by  essential  oils, 
525. 

Rationality  of  birds,  405. 

Rattle-snakes,  difference  of  the 
sexes  in  the,  347;  rattles  as  a 
call,  349. 


INDEX. 


659 


Raven,  vocal  organs  of  the,  366; 
stealing-  bright  objects,  408;  pied, 
of  the  Feroe  Islands,  418. 

Rays,  prehensible  organs  of  male, 
327. 

Razor-bill,  young  of  the,  481. 

Reade,  Winwood,  suicide  among 
savages  in  Africa,  113;  mulat- 
toes  not  prolific,  167;  effect  of 
castration  of  horned  sheep,  502; 
on  the  Guinea  sheep,  230;  on  the 
occurrence  of  a  mane  in  an  Af- 
rican ram,  528;  on  the  negroes' 
appreciation  of  the  beauty  of 
their  women,  572;  on  the  admira- 
tion of  negroes  for  a  black  skin, 
573;  on  the  idea  of  beauty  among 
negroes,  576;  on  the  Jollofs,  582; 
on  the  marriage-customs  of  the 
negroes,  593. 

Reason,  in  animals,  73. 

Redstart,  American,  breeding  in 
immature  plumage,  479. 

Redstarts,  new  mates  found  by, 
403. 

Reduvidse,  stridulation  of,  278. 

Reed-bunting,  head  feathers  of 
the  male,  396;  attacked  by  a 
bullfinch,  407. 

Reefs,  fishes  frequenting,  339. 

Reeks.  H.,  retention  of  horns  by 
breeding  deer,  499;  cow  rejected 
by  a  bull,  520;  destruction  of  pie- 
bald rabbits  by  cats,  537. 

Regeneration,  partial,  of  lost 
parts  in  man,  8. 

Regent-bird,   408. 

Reindeer,  horns  of  the,  228;  bat- 
tles of,  497;  horns  of  the  female, 
499;  antlers  of,  with  numerous 
points,  506;  winter  change  of 
the,  537;  sexual  preferences 
shown  by,  518. 

Relationship,  terms  of.  588. 

Religion,  deficiency  of,  among 
certain  races,  91;  psychical  ele- 
ments of,   92. 

Remorse,  111;  deficiency  of, 
among  savages,   128. 

Rengger,  on  the  diseases  of  Cebus 
Azarse,  7;  on  the  diversity  of 
the  mental  faculties  of  mon- 
keys, 26;  on  the  Payaguas  In- 
dians, 31;  on  the  inferiority  of 
Europeans  to  savages  in  thejr 
senses,  32;  revenge  taken  by 
monkeys,  67;  on  maternal  affec- 
tion in  a  Cebus,  68;  on  the  rea- 
soning powers  of  American 
monkeys,  75;  on  the  use  of 
stones  by  monkeys  for  crack- 
ing hard  nuts,  78;  on  the  sounds 
uttered  by  Cebus  Azarse,  82;  on 
the  signal-cries  of  monkeys,  85; 
on  the  polygamous  habits  of 
Mycetes  caraya,  213;  on  the 
voice  of  the  howling  monkeys, 
523;  on  the  odor  of  Cervus  cam- 
pestris,  524;  on  the  beards  of 
Mycetes   caraya     and     Pithecia 


Satanas,  527;  on  the  colors  of 
Felis  mitls,  530;  on  the  colors  of 
Cervus  paludosus,  532;  on  sexual 
differences  of  color  in  Mycetes, 
532;  on  the  color  of  the  infant 
Guaranys,  553;  on  the  early  ma- 
turity of  the  female  of  Cebus 
azarse,  553;  on  the  beards  of  the 
Guaranys,  556;  on  the  emotional 
notes  employed  by  monkeys,  566; 
on  Am.erican  polygamous  mon- 
keys, 585. 

Representative  species,  of  birds, 
462. 

Reproduction,  unity  of  phenom- 
ena of,  throughout  the  mamma- 
lia,  8;  period  of,   in  birds,   479. 

Reproductive  system,  rudimen- 
tary structures  in  the,  23;  ac- 
cessory parts   of,   156. 

Reptiles,  346. 

and  birds,  alliance  of,  160. 

Resemblances,  small,  between 
man  and  the  apes,  146. 

Retrievers,  exercise  of  reasoning 
faculties  by,  75. 

Revenge,  manifested  by  animals, 
67. 

Reversion,  35;  perhaps  the  cause 
of  some   bad   dispositions,   134. 

Rhagium,  difference  of  color  in 
the  sexes  of  a  species  of,  292. 

Rhamphastos  carinatus,  487. 

Rhea  darwinii,  473. 

Rhinoceros,  nakedness  of,  55; 
horns  of,  501;  horns  of,  used  de- 
fensively, 513;  attacking  white 
or  gray  horses,  535. 

Rhynchaea,  sexes  and  young  of, 
470. 

australis,  470. 

bengalensis,  470. 

capensis,  471. 


Rhythm,  perception  of,  by  ani- 
mals, 564. 

Richard,  M.,  on  rudimentary  mus- 
cles in  man,  12. 

Richardson,  Sir  J.,  on  the  pairing 
of  Tetrao  umbellus,  362;  on  Te- 
trao  urophasianus,  368;  on  the 
drumming  of  grouse,  371;  on  the 
dances  of  Tetra.o  phasianellus, 
376;  on  assemblages  of  grouse, 
400;  on  the  battles  of  male  deer, 
497;  on  the  reindeer,  499;  on  the 
horns  of  the  musk-ox,  501;  on 
antlers  of  the  reindeer  with 
numerous  points,  506;  on  the 
moose,    511. 

,  on    the    Scotch    deerhound, 

512. 

Richter,  Jean  Paul,  on  imagina- 
tion, 72. 

Riedel,  on  profligate  female 
pigeons,  412. 

Riley,  Mr.,  on  mimicry  in  butter- 
flies, 320;  birds'  disgust  at  taste 
of  certain  caterpillars,  322. 

Ring-ouzel,  colors  and  nidification 
of  the,  449. 


660 


INDEX. 


Ripa,  Father,  on  the  difficulty  of 
distinguishing:  the  races  of  the 
Chinese,  163. 

Rivalry,  in  singing,  between  male 
birds,  364. 

River-hog,  African,  tusks  and 
knobs  of  the,  516. 

Rivers,  analogy  of,  to  islands,  154. 

Roach,  brightness  of  male  during 
breeding-season,   336. 

Robbery,  of  strangers,  considered 
honorable,  114. 

Robertson,  Mr.,  remarks  on  the 
development  of  the  horns  in  the 
roebuck  and  red-deer,  229. 

Robin,  pugnacity  of  the  male,  360; 
autumn  song  of  the,  356;  female 
singing  of  the,  365;  attacking 
other  birds  with  red  in  their 
plumage,  407;  young  of  the,  475. 

Robinet,  on  the  difference  of  size 
of  the  male  and  female  cocoons 
of  the  silk-moth,  276. 

Rodents,  uterus  in  the,  36;  ab- 
sence of  secondary  sexual  char- 
acters in,  214;  sexual  differences 
in  the  colors  of,  529. 

Roe,   winter  change  of  the,   537. 

Rohlfs,  Dr.,  Caucasian  features  in 
negro,  163;  fertility  of  mixed 
races  in  Sahara,  167;  colors  of 
birds  in  Sahara,  485;  ideas  of 
beauty  amongst  the  Bornuans, 
576. 

Rolle,  P.,  on  the  origin  of  man, 
2;  on  a  change  in  German  fami- 
lies settled  in  Georgia,  192. 

Roller,  harsh  cry  of,  366. 

Romans,  a,ncient,  gladiatorial  ex- 
hibitions of  the,  119. 

Ronjou,  M.  A.,  coincidence  of  ar- 
rested development  with  poly- 
dactylism,   35. 

Rook,  voice  of  the,  370. 

Rossler,  Dr.,  on  the  resemblance 
of  the  lower  surface  of  butfer- 
fiies  to  the  bark  of  trees,  308. 

Rostrum,  sexual  difference  in  the 
length  of,  in  some  weevils,  204. 

Royer,  Madlle.,    mammals  giving 

Rudimentary  organs,  11;  origin 
of,  23. 

Rudiments,  presence  of,  in  lan- 
guages, 87. 

Rudolphi,  on  the  want  of  connec- 
tion between  climate  and  the 
color  of  the  skin,  187. 

Ruff,  supposed  to  be  polygamous, 
215;  proportion  of  the  sexes  in 
the,  243;  pugnacity  of  the,  357; 
double  moult  in,  3S5,  387;  dura- 
tion of  dances  of,  400;  attraction 
of  the,  to  bright  objects,  407. 

Ruminants,  male,  disappearance 
of  canine  teeth  in,  52,  557;  gen- 
erally polygamous,  213;  suborbit- 
al pits  of,  525:  sexual  differences 
of  color  in,  530. 

Rupicola  crocea,  display  of  plum- 
age by  the  male,  390. 


Ruppell,  on  canine  teeth  in  deer 

and  antelopes,  510. 

Russia,  numerical  proportion  of 
male  and  female  births  in,  211, 
237. 

Ruticilla,  455. 

Rutimeyer,  Prof.,  on  the  physiog- 
nomy of  the  apes,  52;  on  the  sex- 
ual differences  of  monkeys,  556. 

Rutlandshire,  numerical  propor- 
tion of  male  and  female  births 
in,  237. 


Sachs,  Prof.,  on  the  behavior  of 
the  male  and  female  elements,  \n 
fertilization,    218. 

Sacrifices,  human,  140. 

Sagittal  crest  in  male  apes  and 
Australians,     553. 

Sahara,  fertility  of  mixed  races 
in,  167;  birds  of  the,  450;  animal 
inhabitants   of    the,    485. 

Sailors,  growth  of,  delayed  by 
conditions  of  life,  30;  loag- 
sighted,     32. 

Sailors  and  soldiers,  difference  in 
the   proportions    of,   30. 

St.  John,  Mr.,  on  the  attachment 
of   mated    birds,    405. 

St.  Kilda,  beards  of  the  inhabi- 
tants   of,    555. 

Salmo  eriox,  and  S.  umbla,  color- 
ing of  the  male,  during  the 
breeding    season,    336. 

lycaodon,    331. 

salar,   331. 

Salm.on  leaping  out  of  fresh 
water,  104;  male,  ready  to  breed 
before  the  female,  208;  propor- 
tion of  the  sexes  in,  244;  male, 
pugnacity  of  the,  328;  male, 
characters  of  during  the  breed- 
ing season,  330,  336;  spawning  of 
the,  341;  breeding  of  immature 
male,   480. 

Salvin,  O.,  inheritance  of  muti- 
lated feathers,  57,  384,  603;  on  the 
Humming-birds,  380,  597;  on  the 
numerical  proportion  of  the 
sexes  in  Humming-birds,  243, 
483;  on  Chamsepetes  and  Penel- 
ope, 373;  on  Selasphorus  platy- 
cercus,  373;  Pipra  deliciosa, 
374;     on   Chasmorhynchus,    384. 

Samoa  Islands,  beardlessness  of 
the  natives  of,  555,  575. 

Sand-skipper,  266. 

Sandwich  Islands,  variation  in 
the  skulls  of  the  natives  of  the, 
25;  decrease  of  native  popula- 
tion, 181;  population  of,  253;  su- 
periority of  the  nobles  in  the, 
581. 

Islanders,  lice  of,  166. 

San  Giuliano,  women  of,  581. 

Santali,  recent  rapid  increase  of 
the,  44;  Mr.  Hunter  on  the,  187. 


INDEX. 


661 


Saphirina,  characters  of  the 
males  of,  267. 

Sarkidiornis  melanonotus,  char- 
acters of  the  young,  459. 

Sars,  O.,  on  Pontoporeia  afflnis, 
262. 

Saturnia,  carpini.  attraction  of 
males  by  the  female,  248. 

lo,    difference   of  coloration 

in  the  sexes  of  312. 

Saturniidse,  coloration  of  the,  311, 
312. 

Savage,  Dr.,  on  the  fighting  of  the 
male  gorillas,  557;  on  the  habits 
of  the  gorilla,  586. 

and  Wyman  on  the  polyg- 
amous habits  of  the  gorilla,  213. 

Savages,  uniformity  of,  exagger- 
ated, 27;  long-sighted,  32;  rate  of 
increase  among,  usually  small, 
43;  retention  of  the  prehensile 
power  of  the  feet  by,  50;  imita- 
tive faculties  of,  85,  126;  causes 
of  low  morality  of,  115;  tribes  of, 
supplanting  one  another,  125; 
improvements  in  the  arts 
among,  141;  arts  of,  175;  fond- 
ness of  for  rough  music,  375; 
attention  paid  by,  to  personal 
appearance,  568;  relation  of  the 
sexes  among,  586. 

Saviotti,  Dr.,  division  of  malar 
bone,  37. 

Saw-fly,  pugnacity  of  a  male,  289. 

Saw-flies,  proportions  of  the  sexes 
in,  250. 

Saxicola  rubicola,  young  of,  483. 

Scalp,  motion  of  the,  13. 

Scent-glands  in  snakes,  348. 

Schaaffhausen,  Prof.,  on  the  de- 
velopment of  the  posterior  mo- 
lars in  different  races  of  man, 
20;  on  the  jaw  from  La  Naulette, 
39;  on  the  correlation  between 
muscularity  and  prominent  su- 
pra-orbital ridges,  42;  on  the 
mastoid  processes  of  man,  51; 
on  modifications  of  the  cranial 
bones,  54;  on  human  sacrifices, 
140;  on  the  probable  speedy  ex- 
termination of  the  anthropomor- 
phous apes,  152;  on  the  ancient 
inhabitants  of  Europe,  177;  on 
the  effects  of  use  and  disuse 
of  parts,  192;  on  the  superciliary 
ridge  in  man,  551;  on  the  ab- 
sence of  race-differences  in  the 
infant  skull  in  man,  552;  on  ugli- 
ness, 578. 

Schaum,  H.,  on  the  elytra  of 
Dytiscus  and  Hydroporus,  273. 

Schelver,  on  dragon-fiies,  288. 

Schiodte,  on  the  stridulation  of 
Heterocerus,  300. 

Schlegel,  F.  von,  on  the  complexi- 
ty, of  the  languages  of  uncivil- 
ized peoples,  88. 

,  Prof.,  on  Tanysiptera,  463. 

Schleicher,  Prof.,  on  the  origin  of 
language,  84. 
43 


Schomburgk,  Sir  R.,  on  the  pug- 
nacity of  the  male  musk-duck 
of  Guiana,  358;  on  the  courtship 
of  Rupicola  crocea,  390. 

Schoolcraft,  Mr.,  on  the  difiicul- 
ty  of  fashioning-  stone  imple- 
ments, 48. 

Schweinfurth,  complexion  of  ne- 
groes, 551. 

Scieena  aquila,  343. 

Sclater,  P.  L.,  on  modified  sec- 
ondary wing-feathers  in  the 
males  of  Pipra,  373;  on  elongated 
feathers  in  nightjars,  379;  on  the 
species  of  Chasmorhynchus,  384; 
on  the  plumage  of  Pelecanus 
onocrotalus,  388;  on  the  plan- 
tain-eaters, 453;  on  the  sexes  and 
young  of  Tadorna  variegata,  473; 
on  the  colors  of  Lemur  macaco, 
532;  on  the  stripes  in  asses,  543. 

Scolecida,  absence  of  secondary 
sexual  characters  in,   257. 

Scolopax  frenata,  tail-feathers  of, 
372. 

gallinago,  drumming  of,  372. 

javensis,     tail-feathers     of, 


372. 


major,  assemblies  of,  400. 

Wilsonii,      sound    produced 

by,  373. 

Scolytus,  stridulation  of,  299. 

Scoter-duck,  black,  sexual  differ- 
ence in  coloration  of  the,  487; 
bright  beak  of  male,  487. 

Scott,  Dr.  on  idiots  smelling  their 
food,  35. 

,  J.,  on  the  color  of  the  beard 

in  man,  553. 

Scrope,  on  the  pugnacity  of  the 
male  salmon,  330;  on  the  battles 
of  stags,  497. 

Scudder,  S.  H.,  imitation  of  the 
stridulation  of  the  Orthoptera, 
281;  on  the  stridulation  of  the 
Acridiidse,  283;  on  a  Devonian 
insect,   286;  on  stridulation,    561. 

Sculpture,  expression  of  the  ideal 
of  beauty  by,  576. 

Sea-anemones,  bright  colors  of, 
257. 

Sea-bear,  polygamous,  214. 

Sea-elephant,  male,  structure  of 
the  nose  of  the,  524;  polygamous, 
214. 

Sea-lion,  polygamous,  214. 

Seal,  bladder-nose,  523. 

Seals,  their  sentinels  generally  fe- 
males, 97;  evidence  furnished  by, 
on  classification,  145;  polyga- 
mous habits  of,  214;  battles  of 
male,  496;  canine  teeth  of  male, 
498;  sexual  differences,  511;  pair- 
ing of,  518;  sexual  peculiarities 
of,  523;  in  the  coloration  of,  530; 
appreciation  of  music  by,  563. 

Sea-scorpion,  sexual  differences 
in,  334. 

Season,  changes  of  color  in  birds, 
in    accordance    with    the,    384; 


662 


INDEX. 


changes  of  plumage  of  birds  in 
relation  to,  456. 

Seasons,  inheritance  at  corre- 
sponding, 224. 

Sebituani,  African  chief,  trying  to 
alter  a  fashion,  570. 

Sebright  Bantam,  233. 

Secondary  sexual  characters,  203; 
relations  of  polygamy  to,  212; 
transmitted  through  both  sexes, 
221;  gradation  of,  in  birds,  424. 

Sedgwick,  W.,  on  hereditary  ten- 
dency to  produce  twins,  44. 

Seemann,  Dr.,  on  the  different  ap- 
preciation of  music  by  different 
peoples,  564;  on  the  effects  of 
music,  566. 

Seidlitz,  on  horns  of  reindeer,  502. 

Selasphorus  platycercus,  acumin- 
ate first  primary  of  the  male, 
373. 

Selby,  P.  J.,  on  the  habits  of  the 
black  and  red  grouse,  215. 

Selection,  double,  221. 

of  male  by  female  birds,  399, 

416. 

,     methodical,     of     Prussian 

grenadiers,  28. 

sexual,     explanation     of. 


205,  209,  216;  influence  of,  on  the 
coloring  of  Lepidoptera,  317. 
-,     sexual    and  natural,  con- 


trasted, 221. 

Self-command,  habit  of,  inherited, 
112;  estimation  of,  115. 

Self-consciousness,  in  animals,  80. 

Self-preservation,  instinct  of,  108. 

Self-sacrifice,  by  savages,  109;  es- 
timation of  115. 

Semilunar  fold,  17. 

Semnopithecus,  149;   long  hair  on 
the  heads  of  species  of,  146,  598. 

chrysomelas,    sexual   differ- 
ences of  color  in,  533. 

comatus,     ornamental     hair 


on  the  head  of,  544. 

frontatus,    beard,    &c.,    of, 


546. 


nasica,  nose  of,  146. 

nemasus,  coloring  of,  546. 

rubicundus,  ornamental  hair 

on  the  head  of,  543. 

Senses,  inferiority  of  Europeans 
to  savages  in  the,  32. 

Sentinels,  among  animals,  97,  104. 

Serpents,  instinctively  dreaded  by 
apes  and  monkeys,  65,  69. 

Serranus,  hermaphroditism  in, 
157. 

Sex,  inheritance  limited  by,  225. 

Sexes,  relative  proportions  of,  in 
man,  237,  252;  probable  relation 
of  the,  in  primeval  man,  585. 

Sexual  characters,  secondary,  203; 
relations  of  polygamy  to,  212; 
transmitted  through  both  sexes, 
221;  gradation  of,  in  birds,  424. 

and  natural  selection,  con- 
trasted, 221. 


Sexual  characters,  effects  of  the 
loss  of,  226;   limitation  of,   226. 

differences  in  man,  8. 

selection,      explanation     of, 

205,  209,  216;  influence  of,  on  the 
coloring  of  Lepidoptera,  315;  ob- 
jections to,  490;  action  of,  in 
mankind,  589. 

similarity,  251. 


Shaler,  Prof.,  sizes  of  sexes  in 
whales,  511. 

Sharks,  prehensile  organs  of  male, 
327. 

Sharpe,  Dr.,  Europeans  in  the 
tropics,  190. 

,  R.  B.,  on  Tanysiptera  Syl- 
via, 445;  on  Ceryle,  451;  on  the 
young  male  of  Dacelo  Gaudich- 
audi,  461. 

Shaw,  Mr.,  on  the  pugnacity  of 
the  male  salmon,  328. 

,   J.,   on  the    decorations  of 

birds,  377. 

Sheep,  danger-signals  of,  97;  sex- 
ual differences  in  the  horns  of, 
225;  horns  of,  229,  501;  domestic, 
sexual  differences  of,  late  de- 
veloped, 232;  numerical  propor- 
tion of  the  sexes  in,  241;  inheri- 
tance of  horns  by  one  sex,  501; 
effect  of  castration,  502;  mode 
of  fighting  of,  504;  arched  fore- 
heads of  some,  528. 

Merino,  loss  of  horns  in  fe- 
males of,  226;  horns  of,  230. 

Shells,  difference  in  form  of,  in 
male  and  female  Gasteropoda, 
259;  beautiful  colors  and  shapes 
of,  260. 

Shield-drake,  pairing  with  com- 
mon duck,  409;  New  Zealand, 
sexes  and  young  of,  473. 

Shooter,  J.,  on  the  Kaffirs,  574;  on 
the  marriage  customs  of  the 
Kaffirs,  593. 

Shrew  mice,  odor  of,  524. 

Shrike,  Drongo,  454. 

Shrikes,  characters  of  young, 
459. 

Shuckard,  W.  B.,  on  sexual  differ- 
ences in  the  wings  of  Hymenop- 
tera,  274. 

Shyness  of  adorned  male  birds, 
397. 

Siagonium,  proportions  of  the 
sexes  in,  249;  dimorphism  in 
males  of,  296. 

Siam,  proportion  of  male  and  fe- 
male births  in,  239. 

Siamese,  general  beardlessness  of 
the,  555;  notions  of  beauty  of 
the,  573;  hairy  family  of,  596. 

Sidgwick,  H.,  on  morality  in  hy- 
pothetical bee  commLinity,  96; 
our  actions  not  entirely  directed 
by  pain  and  pleasure,  116. 

Siebold,  C.  T.  von,  on  the  pro- 
portion of  sexes  in  the  Apus, 
251;   on   the   auditory  apparatus 


INDEX. 


663 


of    the    stridulant    Orthoptera, 
28L 

Sight,  inheritance  of  long  and 
short,  32. 

Signal-cries  of  monkeys,  85. 

Silk-moth,  proportion  of  the  sexes 
in,  245,  247;  Ailanthus,  prof. 
Canestrini,  on  the  destruction  of 
its  larvae  by  wasps,  247;  differ- 
ence of  size  of  the  male  and  fe- 
male cocoons  of  the,  275;  pair- 
ing of  the,  314. 

Simiadae,  148;  their  origin  and 
divisions,  161. 

SimOarity,  sexual,  220. 

Singing  of  the  Cicadse  and  Ful- 
goridae,  279;  of  tree-frogs,  346; 
of  birds,  object  of  the,  364. 

Sirenia,   nakedness  of,   54. 

Sirex  juvencus,  289. 

Siricidae,  difference  of  the  sexes 
in,  289. 

Siskin,  388;  pairing  with  a  canary, 
410. 

Sitana,  throat-pouch  of  the  males 
of,  350,  353. 

Size,  relative,  of  the  sexes  of  in- 
sects, 275. 

Skin,  movement  of  the,  13;  naked- 
ness of,  in  man,  54;  color  of  the, 
187. 

and  hair,  correlation  of  col- 
or of,  192. 

Skull,  variation  of,  in  man,  25; 
cubic  contents  of,  no  absolute 
test  of  intellect,  52;  Neanderthal, 
capacity  of  the,  53;  causes  of 
modification  of  the,  53;  differ- 
ence of,  in  form  and  capacity, 
in  different  races  of  men,  164; 
variability  of  the  shape  of  the, 
170;  differences  of,  in  the  sexes 
in  man,  552;  artificial  modifica- 
tion of  the  shape  of,  569. 

Skunk,  odor  emitted  by  the,  524; 
white   tail   of,    protective,    538. 

Slavery,  prevalence  of,  114;  of  wo- 
men, 588. 

Slaves,  difference  between  field 
and  house-slaves,  191. 

Sloth,  ornaments  of  male,  529. 

Smell,  sense  of,  in  man  and  ani- 
mals, 17. 

Smith,  Adam,  on  the  basis  of  sym- 
pathy, 103. 

,   Sir  A.,  on  the  recognition 

of  women  by  male  Cynocephali, 
8;  on  revenge  by  a  baboon,  67; 
on  an  instance  of  memory  in  a 
baboon,  72;  on  the  retention  of 
their  color  by  the  Dutch  in 
South  Africa,  188;  on  the  polyg- 
amy of  the  South  African  an- 
telopes, 213;  on  the  polygamy  of 
the  lion,  214;  on  the  proportion 
of  the  sexes  in  Kobus  ellipsip- 
rymnus,  242;  on  Bucephalus  ca- 
pensis,  347;  on  South  African  liz- 
ards, 353;  on  fighting  gnus,  497; 
on  the  horns  of  rhinoceroses, 
501;  on  the  fighting  of  lions,  516; 


on  the  colors  of  the  Cape  Eland, 
531;  on  the  colors  of  the  gnu, 
531;  on  Hottentot  notions  of 
beauty,  573;  disbelief  in  commu- 
nistic  marriages,    5S3. 

Smith,  F.,  on  the  Cynipidss  and 
Tenthredinldae,  250;  on  the  rela- 
tive size  of  the  sexes  of  Acule- 
ate Hymenoptera,  277;  on  the 
difference  between  the  sexes  of 
ants  and  bees,  289;  on  the  stridu- 
lation  of  Trox  sabulosus,  300;  on 
the  stridulation  of  Mononychua 
pseudacori,  302. 

Smynthurus  luteus,  courtship  of. 
277. 

Snakes,  sexual  differences  of, 
347;  mental  powers  of,  348;  male, 
ardency  of,  348. 

"Snarling  muscles,"  39. 

Snipe,  drumming  of  the,  372;  col- 
oration of  the,  486. 

,    painted,   sexes  and  young 

of,  470. 


,  solitary,  assemblies  of,  400. 

Snipes,  arrival  of  male  before  the 
female,  208;  pugnacity  of  male, 
358;  double  moult  in,  385. 

Snow-goose,  whiteness  of  the,  488. 

Social  animals,  affection  of,  for 
each  other,  97;  defense  of,  by  the 
males,  104. 

Sociability,  the  sense  of  duty  con- 
nected with,  95;  impulse  to,  in 
animals,  102;  manifestations  of, 
in  man,  105;  instinct  of,  in  ani- 
mals, 106. 

Sociality,  probable,  of  primeval 
men,  60;  infiuence  of,  on  the  de- 
velopment of  the  intellectual 
faculties,  125;  origin  of,  in  man, 
126. 

Soldiers,  American,  measurements 
of.  31. 

and  sailors,  difference  In  the 

proportions  of,   31. 

Solenostoma,  bright  colors  and 
marsupial  sack  of  the  females 
of,  342. 

Song  of  male  birds  appreciated  by 
their  females,  89;  want  of,  in 
brilliant  plumaged  birds,  395;  of 
birds,  444. 

Sorex,  odor  of,  523. 

Sounds  admired  alike  by  man  and 
animals,  90;  produced  by  fishes, 
343;  produced  by  male  frogs  and 
toads,  346;  instrumentally  pro- 
duced by  birds,  371  et  seq. 

Spain,  decadence  of,  137. 

Sparassus  smaragdulus,  difference 
of  color  in  the  sexes  of,  268. 

Sparrow,  pugnacity  of  the  male, 
356;  acquisition  of  the  Linnet's 
song  by  a,  366;  coloration  of  the, 
467;  immature  plumage  of  the, 
461. 

,    white-crowned,    young    of 

the,  481. 

Sparrows,  house-  and  tree-,  449. 


664 


INDEX. 


Sparrows,  new  mates  found  by, 
403. 

,    sexes    and   young   of,    477; 

learning  to  sing,  565. 

Spathura  Underwoodi,  383. 

Spawning  of  fishes,  337,  340. 

Spear,  used  before  dispersion  of 
man,   175. 

Species,  causes  of  the  advance- 
ment of,  133;  distinctive  charact- 
ers of,  162;  or  races  of  man,  164; 
sterility  and  fertility  of,  when 
crossed,  166;  supposed,  of  man, 
170;  gradation  of,  171;  difficulty 
of  defining,  171;  representative, 
of  birds,  462;  of  birds,  compara- 
tive differences  between  the 
sexes    of    distinct,    463. 

Spectrum  femoratum,  difference 
of  color  in  the  sexes  of,  287. 

Speech,  connection  between  the 
brain  and  the  faculty  of,  85; 
connection  of  intonation  with 
music,  564. 

"Spel"  of  the  black-cock,  370. 

Spencer,  Herbert,  on  the  influence 
of  food  on  the  size  of  the  jaws, 
32;  on  the  dawn  of  intelligence, 
65;  on  the  origin  of  the  belief  in 
spiritual  agencies,  91;  on  the 
origin  of  the  moral  sense,  119; 
on  music,  566. 

Spengel,  disagrees  with  explana- 
tion of  man's  hairlessness,  596. 

Sperm-whales,  battles  of  male,  496. 

Sphingidse,   coloration  of  the,   311. 

Sphinx,  Humming-bird,  313. 

,  Mr.  Bates  on  the  caterpillar 

of  a.   321. 

moth,  musky  odor  of,  305. 


Spiders,  268;  parental  feeling  in, 
102;  male,  more  active  than  fe- 
male, 217;  proportion  of  the 
sexes  in,  250;  secondary  sexual 
characters  of,  268;  courtship  of 
male,  273;  attracted  by  music, 
269;   male,   small   size  of,   269. 

Spilosoma  menthastri,  rejected  by 
turkeys,  320. 

Spine,  alteration  of,  to  suit  the 
erect  attitude  of  man,  51. 

Spirits,  fondness  of  monkeys  for, 
7. 

Spiritual  agencies,  belief  in,  al- 
most universal,  91. 

Splza  cyanea  and  civis,  407. 

Spoonbill,  370;  Chinese,  change  of 
plumage  in,  455. 

Spots,  retained  throughout  groups 
of  birds,  421;  disappearance  of, 
in  adult  mammals,  541. 

Sprengel,  C.  K.,  on  the  sexuality 
of  plants,  208. 

Spring-boc,  horns  of  the,  504. 

Sproat,  Mr.,  on  the  extinction  of 
savages  in  Vancouver  Island, 
178;  on  the  eradication  of  facial 
hair  by  the  natives  of  Van- 
couver Island,  575;  on  the  eradi- 
cation of  the  beard  by  the  In- 
dians of  Vancouver  Island,   598. 


Spurs,  occurrence  of.  In  female 
fowls,  223,  226;  development  of, 
In  various  species  of  Phasiani- 
dse,  230;  of  Gallinaceous  birds, 
358.  360;  development  of,  in  fe- 
male Gallinaceae,  443. 

Squilla,  different  colors  of  the 
sexes  of  a  species  of,  267. 

Squirrels,  battles  of  male,  496;  Af- 
rican, sexual  differences  in  the 
coloring  of,  529;  black,  535. 

Stag,  long  hairs  of  the  throat  of, 
517;  horns  of  the,  222,  224;  battles 
of,  497;  horns  of  the,  with 
numerous  branches,  506;  bellow- 
ing of  the,  521;  crest  of  the,  526. 

beetle,  numerical  proportion 

of  sexes  of,  249;  large  size  of 
male,  276;  weapons  of  the  male, 
296. 

Stainton,  H.  T.,  on  the  numerical 
proportion  of  the  sexes  in  the 
smaller  moths,  247;  habits  of 
Elachista  rufocinerea,  247;  on 
the  coloration  of  moths,  312;  on 
the  rejection  of  Spilosoma  men- 
thastri, by  turkeys,  312;  on  the 
sexes  of  Agrotis  exclamationis, 
313. 

Staley,  Bishop,  mortality  of  in- 
fant Maories,  182. 

Stallion,  mane  of  the,  517. 

Stallions,  two,  attacking  a  third, 
98;  fighting,  497;  small  canine 
teeth  of,  510. 

Stansbury,  Capt.,  observations  on 
pelicans,  99. 

Staphylinidse,  hornlike  processes 
in  male,  296. 

Starfishes,  parental  feeling  in,  102; 
bright  colors  of  some,  257. 

Stark,  Dr.,  on  the  death-rate  in 
towns  and  rural  districts,  135; 
on  the  influence  of  marriage  on 
mortality,  136;  on  the  higher 
mortality  of  males  in  Scotland, 
238. 

Starling,  American  field-,  pugnac- 
ity of  male.  363. 

,   red-winged,    selection  of   a 

mate  by  the  female,  410. 

Starlings,  three,  frequenting  the 
same  nest,  215,  404;  new  mates 
found   by,  403. 

Statues,  Greek,  Egyptian,  Assyri- 
an, &c.,  contrasted,  576. 

Stature,  dependence  of,  upon  local 
influences,  30. 

Staudinger,  Dr.,  on  breeding  Lep- 
idoptera,  247;  his  list  of  Lepi- 
doptera,  248. 

Staunton,  Sir  G.,  hatred  of  in- 
decency a  modern  virtue,   115. 

Stealing  of  bright  objects  by 
birds,  408. 

Stebbing,  T.  R.,  on  the  nakedness 
of  the   human  body,   594. 

Stemmatopus,  524. 

Stendhal,  see  Bombet, 

Stenobothrus  pratorum,  stridula- 
tion,    283. 


INDEX. 


665 


Stephen,  Mr.  L.,  on  the  difference 
in  the  minds  of  men  and  ani- 
mals, 76;  on  general  concepts  in 
animals,  86;  distinction  between 
material  and  formal  morality, 
107. 

Sterility,  general,  of  sole  daugh- 
ters, 132;  when  crossed,  a  dis- 
tinctive character  of  species,  162; 
under  changed  conditions,  184, 
186. 

Sterna,  seasonal  change  of  plu- 
mage in,  488. 

Stickle-back,  polygamous,  216; 
male,  courtship  of  the,  328;  male, 
brilliant  coloring  of,  during  the 
breeding  season,  337;  nidification 
of   the.    341. 

Sticks  used  as  implements  and 
weapons  by  monkeys,  79. 

Sting  in  bees,  204. 

Stokes,  Capt.,  on  the  habits  of  the 
great  bower-bird,  376. 

Stoliczka,  Dr.,  on  colors  in 
snakes,  348. 

Stonechat,  young  of  the,  483. 

Stone  implements,  difficulty  of 
making,  48;  as  traces  of  extinct 
tribes,  177. 

Stones,  used  by  monkeys  for 
breaking  hard  fruits  and  as  mis- 
siles, 47;  piles  of,  175. 

Stork,  black,  sexual  differences  in 
the  bronchi  of  the,  370;  red  beak 
of  the,  487. 

Storks,  487,  489;  sexual  difference 
in  the  color  of  the  eyes  of,  419. 

Strange,  Mr.,  on  the  satin  bower 
bird,  376. 

Stretch,  Mr.,  on  the  numerical 
proportion  in  the  sexes  of  chick- 
ens, 242. 

Strepsiceros  kudu,  horns  of,  508; 
markings  of,  538. 

Stridulation,  by  males  of  Theridi- 
on,  270;  of  the  Orthoptera  and 
Homoptera  discussed,  286;  of 
beetles,  298. 

Stripes,  retained  throughout 
groups  of  birds,  421;  disappear- 
ance of,  in  adult  mammals,  541. 

Strix  flammea,  403. 

Structure,  existence  of  unservice- 
able modifications  of,  59. 

Struggle  for  existence,  in  man,  138, 
142. 

Struthers,  Dr.,  on  the  occurrence 
of  the  supra-condyloid  foramen 
In  the  humerus  of  man,  21. 

Sturnella  ludoviciana,  pugnacity 
of  the  male,  363. 

Sturnus  vulgaris,  403. 

Sub-species,    171. 

Suffering,  in  strangers,  indiffer- 
ence of  savages  to,  114. 

Suicide,  133;  formerly  not  regarded 
as  a  crime,  113;  rarely  practiced 
among  the  lowest  savages,  114. 

Suidse,  stripes  of  young,  458. 


Sulivan,  Sir  B.  J.,  on  speaking  of 
parrots,  83;  on  two  stallions  at- 
tacking a  third,  497. 

Sumatra,  compression  of  the  nose 
by   the  Malays   of,   577. 

Sumner,  Archb.,  man  alone  cap- 
able of  progressive  improve- 
ment, 77. 

Sun-birds,  nidification  of,  448. 

Superstitions,  141;  prevalence  of, 
118. 

Superstitious  customs,  93. 

Superciliary  ridge  in  man,  551,  553. 

Supernumerary  digits,  more  fre- 
quent in  men  than  in  women, 
219;  inheritance  of,  227;  early  de- 
velopment of,  232. 

Supra-condyloid  foramen  in  the 
early  progenitors  of  man,   156. 

Suspicion,  prevalence  of,  among 
animals,  67. 

Swallow-tail  butterfly,   309. 

Swallows  deserting  their  young, 
104,  110. 

Swan,  black,  wild,  trachea  of  the, 
369;  white  young  of,  477;  red 
beak  of  the,  487;  black-necked, 
489. 

Swans,  487,  489;  young,  475. 

Swaysland,  Mr.,  on  the  arrival  of 
migratory  birds,  208. 

Swifts,  migration  of,  104. 

Swinhoe,  R.,  on  the  common  rat 
in  Formosa  and  China,  78;  be- 
havior of  lizards  when  caught, 
351;  on  the  sounds  produced  by 
the  male  hoopce,  371;  on  Dicru- 
rus  macrocercus  and  the  spoon- 
bill, 454;  on  the  young  of  Arde- 
ola,  463;  on  the  habits  of  Tur- 
nix,  471;  on  the  habits  of  Rhyn- 
chaea  bengalensis,  470;  on  Ori- 
oles breeding  in  immature  plu- 
mage, 479. 

Sylvia  atricapilla,  young  of,  482. 

cinerea,  aerial  love-dance  of 

the  male,  376. 

Sympathy,  131;  among  animals, 
99;  its  supposed  basis,  103. 

Sympathies,  gradual  widening  of, 
109. 

Syngnathous  fishes,  abdominal 
pouch  in  male,  158. 

Sypheotides  auritus.  acuminated 
primaries  of  the  male,  373;  ear- 
tufts  of,  379. 


Tabanidse,  habits  of,  204. 

Tadorna  variegata,  sexes  and 
young  of,  473. 

vulpanser,  409. 

Tahitians,  141;  compression  of  the 
nose  by  the,  577. 

Tail,  rudimentary,  occurrence  of, 
in  man,  22;  convoluted  body  in 
the  extremity  of  the,  23;  absence 
of,  in  man  and  the  higher  apes, 


666 


INDEX. 


56;  variability  of,  in  species  of 
Macacus  and  in  baboons,  56; 
presence  of,  in  the  early  progen- 
itors of  man,  156;  length  of,  in 
pheasants,  439,  445,  446;  difference 
of  length  of  the,  in  the  two 
sexes  of  birds,  445. 

Tait,  Lawson,  on  the  effects  of 
natural  selection  on  civilized  na- 
tions, 130. 

Tanager,  scarlet,  variation  in  the 
male,  418. 

Tanagra  sestiva,  age  of  mature 
plumage  in,  478. 

rubra,  418;  young  of,  483. 

Tanais,  absence  of  mouth  in  the 
males  of  some  species  of,  204; 
relations  of  the  sexes  in,  251; 
dimorphic  males  of  a  species  of, 
262. 

Tankerville,  Earl,  on  the  battles 
of  wild  bulls,  497. 

Tanysiptera,  races  of,  determined 
from  adult  males,  463. 

sylvia,  long  tail-feathers  of, 

445. 

Taphroderes  distortus,  enlarged 
left  mandible  of  the  male,  274. 

Tapirs,  longitudinal  stripes  of 
young,  459,  541. 

Tarsi,  dilatation  of  front,  in  male 
beetles,  273. 

Tarsius,  152. 

Tasmania,  half-castes  killed  by 
the  natives  of,  166. 

Tasmanians,   extinction  of,  179. 

Taste,  in  the  Quadrumana,  536. 

Tattooing,  174;  universality  of, 
569. 

Taylor,  G.,  on  Quiscalus  major, 
244. 

Tea,  fondness  of  monkeys  for,  7. 

Tear-sacks,   of  Ruminants,  525. 

Teebay,  Mr.,  on  changes  of  plu- 
mage in  spangled  Hamburgh 
fowls,  224. 

Teeth,  rudimentary  incisor,  in 
Ruminants,  11;  posterior  molar, 
in  man,  19;  wisdom,  20;  diversity 
of,  25;  canine,  in  the  early  pro- 
genitors of  man,  156;  canine,  of 
male  mammals,  498;  in  man,  re- 
duced by  correlation,  557;  stain- 
ing of  the,  569;  front,  knocked 
out  or  filed  by  some  savages, 
569. 

Tegetmeier,  Mr.,  on  the  transmis- 
sion of  colors  in  pigeons  by  one 
sex  alone,  226;  numerical  pro- 
portion of  male  and  female 
births  in  dogs,  240;  on  the 
abundance  of  male  pigeons,  242; 
on  the  wattles  of  game-cocks, 
398;  on  the  courtship  of  fowls, 
411;  on  the  loves  of  pigeons,  412; 
on  dyed  pigeons,  412;  blue  dragon 
pigeons,  440. 

Tembeta,  S.  American  ornament, 
570. 

Temper,  in  dogs  and  horses,  in- 
herited, 67. 


Tench,  proportions  of  the  sexes  in 
the,  245;  brightness  of  male,  dur- 
ing breeding  season,  336. 

TenebrionidEe,  stridulation  of,   299. 

Tennent,  Sir  J.  E.,  on  the  tusks  of 
the  Ceylon  Elephant,  503,  510; 
on  the  frequent  absence  of  beard 
in  the  natives  of  Ceylon,  555;  on 
the  Chinese  opinion  of  the  as- 
pect of  the  Cingalese,  572. 

Tennyson,  A.,  on  the  control  of 
thought,  119. 

Tenthredinidae,  proportions  of 
the  sexes  in,  250;  fighting  habits 
of  male,  289;  difference  of  the 
sexes  in,  289. 

Tephrodornis,  young  of,  462. 

Terai,  in  India,  177. 

Termites,  habits  of,  288. 

Terns,  white,  488;  and  black,  489. 

,  seasonal  change  of  plumage 

in,  488. 

Terror,  common  action  of,  upon 
the  lower  animals  and  man,  67. 

Testudo  elegans,  347. 

nigra,  346. 

Tetrao  cupido,  battles  of,  363;  sex- 
ual difference  in  the  vocal  or- 
gans of,  367. 

phasianellus,  dances  of,  375; 

duration  of  dances  of,  400. 

scoticus,  449,  459,  465. 

tetrix,  449,  459,  465;  pugnacity 

of  the  male,  359. 

umbellus,    pairing    of,    363; 


battles  of,  363;  drumming  of  the 
male,  371. 

urogalloides,  dances  of,  400. 

urogallus,  pugnacity  of  the 


male,  359. 

urophasianus. 


inflation  of 
the  sesophagus  in  the  male,  368. 

Thamnobia,  young  of,  462. 

Thaumalea  picta,  display  of  plu- 
mage by  the  male,  391. 

Thecla,  sexual  differences  of  col- 
oring in  species  of,  307. 

rubi,  protective  coloring  of, 

308. 

Thecophora  fovea,  305. 
Theognis,  selection  in  mankind,  28. 
Theridion,    stridulation    of    males 
of,   270. 

lineatum,  269. 

Thomisus  citreus,  and  T.  flori- 
colens,  difference  of  color  in  the 
sexes  of,  268. 

Thompson,  J.  H.,  on  the  battles 
of  sperm-whales,  496. 

,  W.,  on  the  coloring  of  the 

male  char  during  the  breeding 
season,  337;  on  the  pugnacity  of 
the  males  of  Gallinula  chloro- 
pus,  357;  on  the  finding  of  new 
mates  by  magpies,  402;  on  the 
finding  of  new  mates  by  Pere- 
grine falcons,  402. 

Thorax,  processes  of,  in  male 
beetles,  292. 

Thorell,  T.,  on  the  proportion  of 
the  sexes  in  spiders,  251. 


INDEX. 


667 


Thornback,  difference  in  the  teeth 
of  the  two  sexes  of  the,  331. 

Thoug-hts,  control  of,  119. 

Thrush,  pairing-  with  a  blackbird, 
409;  colors  and  nidification  of 
the,  449. 

Thrushes,  characters  of  young-, 
449,   459. 

Thug",  remorse  of  a,   114. 

Thumb,  absence  of,  in  Ateles  and 
Hylobates,  49. 

Thury,  M.,  on  the  numerical  pro- 
portion of  male  and  female 
births  among  the  Jews,  238. 

Thylacinus,  possession  of  the 
marsupial  sack  by  the  male,  157. 

Thysanura,  277. 

Tibia,  dilated,  of  the  male  Crabro 
cribrarius,  273. 

T and  femur,  proportions  of,  in 

the  Aymara  Indians,  33. 

Tierra  del  Fuego,  marriage-cus- 
toms of,  592. 

Tig-er,  colors  and  markings  of  the, 
540. 

Tigers,  depopulation  of  districts 
by,  in  India,  44. 

Tillus  el®ngatus,  difference  of 
color  in  the  sexes  of,  291. 

Timidity,  variability  of,  in  the 
same  species,  67. 

Tincae,  proportion  of  the  sexes  in, 
245. 

Tipulse,  pugnacity  of  male,  278. 

Tits,  sexual  difference  of  color  in, 
451. 

Toads,  345;  male,  treatment  of  ova 
by  some,  158;  male,  ready  to 
breed  before  the  female,  208. 

Todas,  infanticide  and  proportion 
of  sexes,  252;  practice  polyandry, 
588;  choice  of  husbands  amongst, 
588. 

Toe,  great,  condition  of,  in  the 
human  embryo,  11. 

Tomicus  villosus,  proportion  of 
the  sexes  in,  250. 

Tomtit,  blue,  sexual  difference  of 
color  in  the,  451. 

Tonga  Islands,  beardlessness  of 
the  natives  of,  555,  575. 

Tooke,  Home,  on  language,  83. 

Tools,  flint,  141;  used  by  monkeys, 
79;  use  of,  47. 

Topknots  in  birds,  377. 

Tortoise,    voice   of   the  male,    561. 

Tortures,  submitted  to  by  Amer- 
ican savages,  115. 

Totanus,  double  moult  in,  385. 

Toucans,  colors  and  nidification 
of  the,  449;  beaks  and  ceres  of 
the,  487. 

Towns,  residence  in,  a  cause  of 
diminished  stature,  30. 

Toynbee,  J.,  on  the  external  shell 
of  the  ear  in  man,  14. 

Trachea,  convoluted  and  imbed- 
ded in  the  sternum,  in  some 
birds,  369;  structure  of  the,  in 
Rhynchsea,  470. 


Trades,  affecting  the  form  of  the 
skull,  54. 

Tragelaphus,  sexual  differences  of 
color  in,  530. 

scriptus,  dorsal  crest  of,  526; 

markings  of,   538,   539. 

Tragopan,  215;  swelling  of  the 
wattles  of  the  male,  during 
courtship,  377;  display  of  plu- 
mage by  the  male,  391;  markings 
of  the  sexes  of  the,  422. 

Tragops  dispar,  sexual  difference 
in  the  color  of,  347. 

Training,  effect  of,  on  the  mental 
difference  between  the  sexes  of 
man,  560. 

Transfer  of  male  characters  to  fe- 
male  birds,   465. 

Transmission,  equal,  of  ornament- 
al characters,  to  both  sexes  in 
mamm-als,    537. 

Traps,  avoidance  of,  by  animals, 
77;  use  of,  47. 

Treachery,  to  comrades,  avoid- 
ance of,  by  savages,  107. 

Tremex   columbEe,   289. 

Tribes,  extinct,  125;  extinction  of, 
178. 

Trichius,  difference  of  color  in  the 
sexes  of  a  species  of,  291. 

Trigla,    343. 

Trimen,  R.,  on  the  proportion  of 
the  sexes  in  South  African  but- 
terflies, 246;  on  the  attraction  of 
males  by  the  female  of  Lasio- 
campa  quercus,  248;  on  Pneu- 
mora,  284;  on  difference  of  color 
in  the  sexes  of  beetles,  291;  on 
moths  brilliantly  colored  be- 
neath, 312;  on  mimicry  in  butter- 
flies, 321;  on  Gynanisa  Isis,  and 
on  the  ocellated  spots  of  Lepi- 
doptera,  422;  on  Cyllo  Leda,  423. 

Tringa,  sexes  and  young  of,  480. 

canutus,  386. 

Triphsena,  coloration  of  the  spe- 
cies of,  310. 

Tristram,  H.  B.,  on  unhealthy  dis- 
tricts in  North  Africa,  189;  on 
the  habits  of  the  chaffinch  in 
Palestine,  244;  on  the  birds  of 
the  Sahara,  450;  on  the  animals 
inhabiting  the  Sahara,  485. 

Triton  cristatus,  344. 

palmipes,  344. 

punctatus,  345. 

Troglodyte     skulls,   greater    than 

those  of  modern  Frenchmen,  53. 
Troglodytes  vulgaris,   467. 
Trogons,  colors  and  nidification  of 

the,   449,   450. 
Tropic-birds,     white    only     when 

mature,  488. 
Tropics,  freshwater  fishes  of  the, 

399. 
Trout,  proportion  of  the  sexes  in, 

245;  male,  pugnacity  of  the,  328, 
Trox   sabulosus,     stridulation   of, 

300. 


668 


INDESX. 


Truth,  not  rare  between  members 
of  the  same  tribe,  114;  more 
highly  appreciated  by  certain 
tribes,  118. 

Tulloch,  Major,  on  the  immunity 
of  the  negro  from  certain  fe- 
vers, 189. 

Tumbler,  almond,  change  of  plu- 
mage in  the,  233. 

Turdus  merula,  449;  young  of,  487. 

migratorius,  459. 

miisicus,  449. 

polyglottus,  young  of,  482. 

■ torquatus,  449^. 

Turkey,  wild,  pugnacity  of  young 
male,  362;  wild,  notes  of  the,  370; 
swelling  of  the  wattles  of  the 
male,  377;  variety  of,  with  a  top- 
knot, 379;  recognition  of  a  dog 
by  a,  407;  male,  wild,  acceptable 
to  domesticated  females,  418; 
wild,  first  advances  made  by 
older  females,  414;  wild,  breast- 
tuft  of  bristles  of  the,  455. 

Turkey-cock,  scraping  of  the 
wings  of,  upon  the  ground,  371; 
wild,  display  of  plumage  by,  390; 
fighting  habits  of,  398. 

Turner,  Prof.  W.,  on  muscular 
fasciculi  in  man  referable  to  the 
panniculus  carnosus,  13;  on  the 
occurrence  of  the  supra-condy- 
loid  foramen  in  the  human  hu- 
merus, 21;  on  muscles  attached 
to  the  coccyx  in  man,  22;  on  the 
filum  terminale  in  man,  22;  on 
the  variability  of  the  muscles, 
26;  on  abnormal  conditions  of 
the  human  uterus,  37;  on  the  de- 
velopment of  the  mammary 
glands,  157;  on  male  fishes  hatch- 
ing ova  in  their  mouths,  158,  341; 
on  the  external  perpendicular 
fissure  of  the  brain,  195;  on  the 
bridging  convolutions  in  the 
brain  of  a  chimpanzee,  196. 

Turnix,  sexes  of  some  species  of, 
470,  474. 

Turtle-dove,   cooing  of  the,  370. 

Tuttle,  H.,  on  the  number  of  spe- 
cies of  man,  170. 

Tylor,  E.  B.,  on  emotional  cries, 
gestures,  &c.,  of  man,  82;  on  the 
origin  of  the  belief  in  spiritual 
agencies,  91;  remorse  for  viola- 
tion of  tribal  usage  in  marrying, 
112;  on  the  primitive  barbarism 
of  civilized  nations,  140;  on  the 
origin  of  counting,  140;  inven- 
tions of  savages,  141;  on  re- 
semblances, of  the  mental  char- 
acters in  different  races  of  man, 
174. 

Type  of  structure,  prevalence  of, 
159. 

TyphcBus,   stridulating  organs  of, 

299;  stridulation  of,  300. 
Twins,      tendency      to      produce, 
hereditary,  44. 


Ugliness,  said  to  consist  in  an  ap- 
proach to  the  lower  animals,  578. 

Umbrella-bird,  368. 

Umbrina,  sounds  produced  by,  343. 

United  States,  rate  of  increase  in, 
43;  influence  of  natural  selection 
on  the  progress  of,  138;  change 
undergone  by  Europeans  in  the, 
191. 

Upupa  epops,  sounds  produced  by 

the  male,  371. 
Uraniidse,  coloration  of  the,  311. 
Uria  troile,  variety  of  (equals  U. 

1  aery  mans),  418. 
Urodela,  344. 

Urosticte  Benjamini,  sexual  dif- 
ferences in,  486. 

Use  and  disuse  of  parts,  effects  of, 
31;  influence  of,  on  the  races  of 
man,  192. 

Uterus,  reversion  in  the,  36;  more 
or  less  divided,  in  the  human 
subject,  37,  42;  double,  in  the 
early  progenitors  of  man,  156. 


Vaccination,  influence  of,  ISO. 

Vancouver  Island,  Mr.  Sproat  on 
the  savages  of,  178;  natives  of, 
eradication  of  facial  hair  by  the, 
575. 

Vanellus  cristatus,  wing  tubercles 
of  the  male,  361. 

Vanessse,  305;  resemblance  of 
lower  surface  of,  to  bark  of 
trees,  308. 

Variability,  causes  of,  27;  in  man, 
analogous  to  that  in  the  lower 
animals,  28;  of  the  races  of  man, 
170;  greater  in  men  than  in 
women,  218;  period  of,  relation 
of  the,  to  sexual  selection,  235; 
of  birds,  416;  of  secondary  sex- 
ual characters  in  man,  554. 

Variation,  laws  of,  28;  correlated, 
41;  in  man,  142;  analogous,  148; 
analogous,  in  plumage  of  birds, 
380. 

Variations,  spontaneous,  42. 

Varieties,  absence  of,  between 
two  species,  evidence  of  their 
distinctness,  163. 

Variety,  an  object  in  nature,  488. 

Variola,  communicable  between 
man  and  the  lower  animals,  7. 

Vaureal,  human  bones  from,  22. 

Veddahs,  monogamous  habits  of, 
586. 

Veitch,  Mr.,  on  the  aversion  of 
Japanese  ladies  to  whiskers,  575. 


INDEX. 


669 


Vengeance,  instinct  of,  110. 

"Venus  Erycina,  priestesses  of,  581. 

Vermes,  261. 

Vermiform  appendage,  21. 

Verreaux,  M.,  on  the  attraction  of 
numerous  males  by  the  female 
of  an  Australian  Bombyx,  248. 

Vertebrae,  caudal,  number  of,  in 
macaques  and  baboons,  56;  of 
monkeys,  partly  imbedded  in 
the  body,  57. 

Vertebrata,  327;  common  origin  of 
the,  153;  most  ancient  progeni- 
tors of,  156;  origin  of  the  voice 
in  air-breathing,  561. 

Vesicula  prostatica,  the  homo- 
logue  of  the  uterus,  23,  157. 

Vibrissae,  represented  by  long 
hairs  in  the  eyebrows,  18. 

Vidua,  397,  456. 

axillaris,   215. 

Villerme,  M.,  on  the  influence  of 
plenty  upon  stature,  30. 

Vinson,  Aug.,  courtship  of  male 
spider,  269;  on  the  male  of  Ep- 
eira  nigra,  269. 

Viper,  difference  of  the  sexes  in 
the,  347. 

Virey,  on  the  number  of  species 
of  man,  170. 

Virtues,  originally  social  only,  113; 
gradual  appreciation  of,   128. 

Viscera,  variability  of,  in  man,  26. 

Vlacovich,  Prof.,  on  the  ischio- 
pubic  muscle,  40. 

Vocal  music  of  birds,  364. 

. organs  of  man,  85;  of  birds, 

87,  444;  of  frogs,  346;  of  the  Inses- 
sores,  366;  difference  of,  in  the 
sexes  of  birds,  367;  primarily 
used  in  relation  to  the  propaga- 
tion of  the  species,  561, 

Vogt,  Karl,  on  the  origin  of  spe- 
cies, 1;  on  the  origin  of  man,  3; 
on  the  semilunar  fold  in  man, 
17;  on  microcephalous  idiots,  34; 
on  the  imitative  faculties  of  mi- 
crocephalous idiots,  85;  on  skulls 
from  Brazilian  caves,  164;  on  the 
evolution  of  the  races  of  man, 
173;  on  the  formation  of  the 
skull  in  women,  552;  on  the 
Ainos  and  negroes,  555;  on  the 
increased  cranial  difference  of 
the  sexes  in  man  with  race  de- 
velopment, 560;  on  the  obliquity 
of  the  eye  in  the  Chinese  and 
Japanese,  572. 

Voice  in  mammals,  521;  in  mon- 
keys and  man,  553;  in  man,  561; 
origin  of,  in  air-breathing  verte- 
brates, 561. 

Von  Baer,  definition  of  advance- 
ment in  the  organic  scale,  159. 

Vulpian,  Prof.,  on  the  resem- 
blance between  the  brains  of 
man  and  of  the  higher  apes,  6. 

Vultures,  selection  of  a  mate  by 
the  female,  410;  colors  of,  488. 


W 

Waders,  young  of,  481. 

Wagner,  R.,  on  the  occurrence  of 
the  diastema  in  a  Kaffir  skull, 
39;  on  the  bronchi  of  the  black 
stork,    370. 

Wagtail,  Ray's,  arrival  of  the 
male  before  the  female,  208. 

Wagtails,  Indian,  young  of,  463. 

Waist,  proportions  of,  in  soldiers 
and  sailors,  31. 

Waitz,  Prof.,  on  the  number  of 
species  of  man,  170;  on  the  lia- 
bility of  negroes  to  tropical  fe- 
vers after  residence  in  a  cold 
climate,  189;  on  the  color  of  Aus- 
tralian infants,  553;  on  the 
beardlessness  of  negroes,  555;  on 
the  fondness  of  mankind  for 
ornaments,  568;  on  negro  ideas 
of  female  beauty,  573;  on  Javan 
and  Cochin  Chine'se  ideas  of 
beauty,   574. 

Walckenaer  and  Gervais,  spider 
attracted  by  music,  270;  on  the 
Myriapoda,  271. 

Waldeyer,  M.,  on  the  hermaph- 
roditism of  the  vertebrate  em- 
bryo, 157. 

Wales,  North,  numerical  propor- 
tion of  male  and  female  births 
in.   237. 

Walker,  Alex.,  on  the  large  size 
of  the  hands  of  laborers'  chil- 
dren, 32. 

,  F.,  on  sexual  differences  in 

the  diptera,  277. 

Wallace,  Dr.  A.,  on  the  prehensile 
use  of  the  tarsi  in  male  moths, 
205;  on  the  rearing  of  the  Ailan- 
thus  silkmoth,  247;  on  breeding 
Lepidoptera,  247;  proportion  of 
sexes  of  Bombyx  cynthia,  B. 
yamamai,  and  B.  Pernyi  reared 
by,  249;  on  the  development  of 
Bombyx  cynthia  and  B.  yam- 
amai, 276;  on  the  pairing  of 
Bombyx  cynthia,  314. 

,    A.    R.,     on   the    origin    of 

man,  3;  on  the  power  of  imita- 
tion in  man,  66;  on  the  use  of 
missiles  by  the  orang,  79;  on  the 
varying  appreciation  of  truth 
among  different  tribes,  118;  on 
the  limits  of  natural  selection 
in  man,  47,  124;  on  the  occur- 
rence of  remorse  among  sav- 
ages, 128;  on  the  effects  of  nat- 
ural selection  on  civilized  na- 
tions, 130;  on  the  use  of  the  con- 
vergence of  the  hair  at  the  el- 
bow in  the  orang,  147;  on  the 
contrast  in  the  characters  of  the 
Malays  and  Papuans,  164;  on  the 
line  of  separation  between  the 
Papuans  and  Malays,  165;  on  the 
birds  of  paradise,  214;  on  the 
sexes  of  Ornithoptera  Croesus, 
246;  on  protective  resemblances, 


670 


INDEX. 


258;  on  the  relative  sizes  of  the 
sexes  of  insects,  276;  on  Ela- 
phomyia,  277;  on  the  pugnacity 
of  the  males  of  Leptorhynchus 
angustatus,  296;  on  sounds,  pro- 
duced by  Euchirus  longima'nus, 
301;  on  the  colors  of  Diadema, 
306;  on  Kallima,  308;  on  the  pro- 
tective coloring  of  moths,  310;  on 
bright  coloration  as  protective 
in  butterflies,  311;  on  variability 
in  the  Papilionidse,  316;  on  male 
and  female  butterflies  inhabit- 
ing different  stations,  317;  on  the 
protective  nature  of  the  dull 
coloring  of  female  butterflies, 
317  318,  321;  on  mimicry  in  but- 
terflies, 320;  on  the  bright  colors 
of  caterpillars,  321;  on  brightly- 
colored  flshes  frequenting  reefs, 
339;  on  the  coral  snakes,  349;  on 
Paradisea  apoda,  380;  on  the  dis- 
play of  plumage  by  male  birds 
of  paradise,  390;  on  assemblies 
of  birds  of  paradise,  400;  on  the 
instability  of  the  ocellated  spots 
in  Hipparchia  Janira,  422;  on 
sexually  limited  inheritance,  438; 
on  the  sexual  coloration  of 
birds,  446,  466,  467,  469,  474;  on  the 
relation  between  the  colors  and 
nidiflcation  of  birds,  446,  449;  on 
the  coloration  of  the  Cotingidse, 
453-  on  the  females  of  Paradisea 
apoda  and  papuana,  464;  on  the 
incubation  of  the  cassowary,  472; 
on  protective  coloration  in  birds, 
485;  on  the  Babirusa,  515;  on  the 
markings  of  the  tiger,  540;  on  the 
beards  of  the  Papuans,  555;  on 
the  hair  of  the  Papuans,  569;  on 
the  distribution  of  hair  on  the 
human  body,  594. 

Walrus,  development  of  the  nicti- 
tating membrane  in  the,  17; 
tusks  of  the,  498,  503;  use  of  the 
tusks  by  the,  509. 

Walsh,  B.  D.,  on  the  proportion 
of  the  sexes  in  Papilio  Turnus, 
254;  on  the  Cynipidse  and  Ceci- 
domyidse,  250;  on  the  jaws  of 
Ammophila,  272;  on  Corydalis 
cornutus,  272;  on  the  prehensile 
organs  of  male  insects,  272;  on 
the  antennge  of  Penthe,  273;  on 
the  caudal  appendages  of  drag- 
on-flies, 274;  on  Platyphyllum 
concavum,  283;  on  the  sexes  of 
the  Ephemeridse,  287;  on  the 
difference  of  color  in  the  sexes 
of  Spectrum  femoratum,  287;  on 
sexes  of  aragon-flies,  288;  on  the 
difference  of  the  sexes  in  the 
Ichneumonidse,  289;  on  the 
sexes  of  Crsodacna  atra,  291;  on 
the  variation  of  the  horns  of  the 
male  Phanaeus  carnifex,  294;  on 
the  coloration  of  the  species  of 
Anthocharis,  309. 


Wapiti,  battles  of,  497;  traces  of 
horns  in  the  female,  500;  attack- 
ing a  man,  506;  crest  of  the 
male,  528;  sexual  difference  in 
the  color  of  the,  531. 

Warbler,  hedge-,  467;  young  of 
the,   475. 

Warblers,  superb,  nidiflcation  of. 
448. 

Wariness,  acquired  by  animals,  77. 

Warington,  R.,  on  the  habits  of 
the  stickle-backs,  328,  341;  on  the 
brilliant  colors  of  the  male 
stickle-back  during  the  breeding 
season,  337. 

Wart-hog,  tusks  and  pads  of  the, 
515. 

Watchmakers,  short-sighted,  32. 

Waterhen,  356. 

Waterhouse,  C.  O.,  on  blind  bee- 
tles, 291;  on  difference  of  color 
in  the  sexes  of  beetles,  291. 

,  G.  R.,  on  the  voice  of  Hylo- 

bates  agilis,  562. 

Water-ouzel,  449;  autumn  song  of 
the,  365. 

Waterton,  C,  on  the  Bell-bird, 
384;  on  the  pairing  of  a  Canada 
goose  with  a  Bernicle  gander, 
409;  on  hares  flghting,  496. 

Wattles,  disadvantageous  to  male 
birds  in  flghting,  398. 

Weale,  J.  Mansel,  on  a  South  Af- 
rican caterpillar,  322. 

Wealth,  influence  of,  131. 

Weapons,  used  by  man,  47;  em- 
ployed by  monkeys,  79;  offensive, 
of  males,  206;  of  mammals,  497 
et  seq. 

Weaver-bird,  366. 

Weaver-birds,  rattling  of  the 
wings  of,  371;  assemblies  of,  400. 

Webb,  Dr.,  on  the  wisdom  teeth, 
20. 

Wedderburn,  Mr.,  assembly  of 
black  game,  401. 

Wedgwood,  Hensleigh,  on  the  ori- 
gin of  language,  84. 

Weevils,  sexual  difference  in 
length  of  snout  in  some,  204. 

Weir,  Harrison,  on  the  numerical 
proportion  of  the  sexes  in  pigs 
and  rabbits,  241;  on  the  sexes  of 
young  pigeons,  242;  on  the  songs 
of  birds,  364;  on  pigeons,  406;  on 
the  dislike  of  blue  pigeons  to 
other  colored  varieties,  412;  on 
the  desertion  of  their  mates  by 
female  pigeons,  412. 

J.  Jenner,  on  the  nightin- 
gale and  blackcap,  208;  on  the 
relative  sexual  maturity  of  male 
birds,  209;  on  female  pigeons  de- 
serting a  feeble  mate,  210;  on 
three  starlings  frequenting  the 
same  nest,  215;  on  the  proportion 
of  the  sexes  in  Machetes  pug- 
nax  and  other  birds,  242,  243;  on 
the  coloration  of  the  Tri- 
phEense,  310;    on  the  rejection  of 


INDEX. 


671 


certain  caterpillars  by  birds,  322; 
on  sexual  differences  of  the  beak 
in  the  goldfinch,  356;  on  a  piping- 
bullfinch,  365;  on  the  object  of 
the  nightingale's  song,  364;  on 
songbirds,  365;  on  the  pugnacity 
of  male  fine-plumaged  birds, 
395;  on  the  courtship  of  birds, 
396;  on  the  finding  of  new  mates 
by  Peregrine-falcons  and  Kes- 
trels, 402;  on  the  bullfinch  and 
starling,  403;  on  the  cause  of 
birds  remaining  unpaired,  409; 
on  starlings  and  parrots  living 
in  triplets,  404;  on  recognition  of 
color  by  birds,  407;  on  hybrid 
birds,  409;  on  the  selection  of  a 
greenfinch  by  a  female  canary, 
410;  on  a  case  of  rivalry  of  fe- 
male bullfinches,  414;  on  the  ma- 
turity of  the  golden-pheasant, 
478. 

Weisbach,  Dr.,  measurement  of 
men  of  diiferent  races,  163;  on 
the  greater  variability  of  men 
than  of  women,  218;  on  the  rela- 
tive proportions  of  the  body  in 
the  sexes  of  different  races  of 
man,  554. 

Weismann,  Prof.,  colors  of  Lyooe- 
nse,  309. 

Welcker,  M.,  on  brachycephaly 
and  dolichocephaly,  54;  on  sex- 
ual differences  in  the  skull  in 
man,  552. 

Wells,  Dr.,  on  the  immunity  of 
colored  races  from  certain  pois- 
ons, 188. 

Westring,  on  the  stridulation  of 
males  of  Theridion,  270;  on  the 
stridulation  of  Reduvius  person- 
atus,  279;  on  the  stridulation  of 
beetles,  300;  on  the  stridulation 
of  Omaloplia  brunnea,  300;  on 
the  stridulating  organs  of  the 
Coleoptera,  301;  on  sounds  pro- 
duced by  Cychrus,   301. 

Westropp,  H.  M.,  on  reason  in  a 
bear,  74;  on  the  prevalence  of 
certain  forms  of  ornamentation, 
175. 

Westwood,  J.  O.,  on  the  classifica- 
tion of  the  Hymenoptera,  144;  on 
the  Culicidas  and  Tabanidee,  204; 
on  a  Hymenopterous  parasite 
with  a  sedentary  male,  217;  on 
the  proportions  of  the  sexes  in 
Lucanus  cervus  and  Siagonium, 
249;  on  the  absence  of  ocelli  in 
female  mutillidae,  272;  on  the 
jaws  of  Ammophila,  273;  on  the 
copulation  of  insects  of  distinct 
species,  272;  on  the  male  of 
Crabro  cribrarius,  273;  on  the 
pugnacity  of  male  Tipulse,  278; 
on  the  stridulation  of  Pirates 
stridulus,  281;  on  the  Cicadae, 
279;  on  the  stridulating  organs 
of  the  crickets,  281;  on  Ephip- 
piger  vitium,  282,  284;  on  Pneu- 
mora,  284;  on  the  pugnacity  of 


the  Mantides,  286;  on  Platyblem- 
nus,  287;  on  difference  in  the 
sexes  of  the  Agrionidse,  287;  on 
the  pugnacity  of  the  males  of  a 
species  of  Tenthredinse,  289;  on 
the  pugnacity  of  the  male  stag- 
beetle,  297;  on  Bledius  taurus 
and  Siagonium,  296;  on  lamelli- 
corn  beetles,  298;  on  the  colora- 
tion of  Lithosia,  311. 

Whale,  Sperm-,  battles  of  male, 
496. 

Whales,  nakedness  of,  54. 

Whately,  Archb.,  language  not 
peculiar  to  man,  82;  on  the  prim- 
itive civilization  of  man,  140. 

Whewell,  Prof.,  on  maternal  af- 
fection, 68. 

Whiskers,  in  monkeys,  146. 

White,  P.  B.,  noise  produced  by 
Hylophila,  305. 

,   Gilbert,    on  the  proportion 

of  the  sexes  in  the  partridge, 
243;  on  the  house-cricket,  280;  on 
the  object  of  the  song  of  birds, 
364;  on  the  finding  of  new  mates 
by  white  owls,  403;  on  spring 
coveys  of  male  partridges,  404. 

Whiteness,  a  sexual  ornament  in 
some  birds,  490;  of  mammals  in- 
habiting snowy  countries,  537. 

White-throat,  aerial  love-dance  of 
the  male,  376. 

Whitney,  Prof.,  on  the  develop- 
ment of  language,  83;  language 
not  indispensable  for  thought, 
86. 

Widow-bird,  polygamous,  214; 
breeding  plumage  of  the  male, 
387,  397;  female,  rejecting  the  un- 
adorned male,   413. 

Widows  and  widowers,  mortality 
of,  136. 

Widgeon,  pairing  with  a  pintail 
duck,  409. 

Wilckens,  Dr.,  on  the  modification 
of  domestic  animals  in  moun- 
tainous regions,  34;  on  a  numer- 
ical relation  between  the  hairs 
and  excretory  pores  in  sheep, 
193. 

Wilder,  Dr.  Burt,  on  the  greater 
frequency  of  supernumerary 
digits  in  men  than  in  women, 
219. 

Williams,  on  the  marriage-cus- 
toms of  the  Fijians,  593. 

Wilson,  Dr.,  on  the  conical  heads 
of  the  natives  of  North-Western 
America,  577;  on  the  Fijians,  577; 
on  the  persistence  of  the  fashion 
of   compressing  the  skull,  578. 

Wing-spurs,  443. 

Wings,  differences  of,  in  the  two 
sexes  of  butterflies  and  Hymen- 
optera, 274;  play  of,  in  the  court- 
ship of  birds,  395. 

Winter,  change  of  color  of  mam- 
mals  in.   537. 

Witchcraft,  93. 


672 


INDEX. 


Wives,  traces  of  the  forcible  cap- 
ture of,  140. 

Wolf,  winter  change  of  the,  537. 

Wolff,  on  the  variability  of  the 
viscera  in  man,  26. 

Woliaston,  T.  V.,  on  Eurygna- 
thus,  274;  on  musical  Curculion- 
idse,  301;  on  the  stridulation  of 
Acalles,  303. 

Wolves  learning  to  bark  from 
dogs,  70;  hunting  in  packs,  98. 

,  black,  535. 

Wombat,  black  varieties  of  the, 
535. 

Women  distinguished  from  men 
by  male  monkeys,  8;  preponder- 
ance of,  in  numbers,  238;  selec- 
tion of,  for  beauty,  391;  effects 
of  selection  of,  in  accordance 
with  different  standards  of 
beauty,  579;  practice  of  captur- 
ing, 584,  587;  early  betrothals  and 
slavery  of,  588;  freedom  of  selec- 
tion by,   in  savage  tribes,  593. 

Wonder,  manifestations  of,  by  an- 
imals,   69. 

Wonfor,  Mr.,  on  sexual  peculiari- 
ties in  the  wings  of  butterflies, 
275. 

Wood,  J.,  on  muscular  variations 
in  man,  26,  40,  41;  on  the  greater 
variability  of  the  muscles  in 
men  than  in  women,  219. 

,   T.   W.,    on  the  coloring  of 

the  orange-tip  butterfly,  310;  on 
the  habits  of  the  Saturniidse, 
312;  quarrels  of  chamaeleons,  353; 
on  the  habits  of  Menura  Al- 
berti,  366;  on  Tetrao  cupido,  368; 
on  the  display  of  plumage  by 
male  pheasants,  391;  on  the  ocel- 
lated  spots  of  the  Argus  pheas- 
ant, 484;  on  the  habits  of  the  fe- 
male cassowary,  472. 

Woodcock,   coloration  of  the,  486. 

Woodpecker,  selection  of  a  mate 
by  the  female,  410. 

Woodpeckers,  366;  tapping  of,  371; 
colors  and  nidiflcation  of  the, 
449,  452,  485;  characters  of  young, 
459,  468,  475. 

Woolner,  Mr.,  observations  on  the 
ear  in  man,  15. 

Wormald,  Mr.,  on  the  coloration 
of  Hypopyra,  312. 

Wounds,  healing  of,  8. 

Wren,  467;  young  of  the,  475. 

Wright,  C,  A.,  on  the  young  of 
Orocetes  and  Petrocincla,  482. 

,  Chauncey,  great  brain- 
power requisite  for  language,  47; 
on  correlative  acquisition,  565; 
on  the  enlargement  of  the  brain 
in  man,  604. 

Mr.,    on   the    Scotch    deer- 


hound,  512;  on  sexual  preference 
in  dogs,  519;  on  the  rejection  of 
a  horse  by  a  mare,  520. 

-,   W.   von,  on  the  protective 


plumage  of  the  Ptarmigan,  385. 
.Writing,  140.  C    I    Q  I    v 


Wyman,  Prof.,  on  the  prolonga* 
tion  of  the  coccyx  in  the  human 
embryo,  11;  on  the  condition  of 
the  great  toe  in  the  human  em- 
bryo, 11;  on  the  occurrence  of 
the  supra-condyloid  foramen  in 
the  humerus  of  man,  21;  on  vari- 
ation in  the  skulls  of  the  natives 
of  the  Sandwich  Islands,  25;  on 
the  hatching  of  the  eggs  in  the 
mouths  and  branchial  cavities 
of  male  fishes,  158,  341. 


Xenarchus,    on   the  Cicadse,   279. 
Xenophon,     selection  in  mankind 

advocated  by,  28. 
Xenorhynchus,   sexual    difference 

in  the  color  of  the  eyes  in,  419. 
Xiphophorus       Hellerii,     peculiar 

anal  fin  of  the  male,  334. 
Xylocopa,  difference  of  the  sexes 

in,  290. 


Tarrell,  W.,  on  the  habits  of  the 
Cyprinidse,  245;  on  Raia  clavata, 
327;  on  the  characters  of  the 
male  salmon  during  the  breed- 
ing season,  330,  366;  on  the  char- 
acters of  the  rays,  327,  331;  on 
the  gemmeous  dragonet,  333;  on 
the  spawning  of  the  salmon,  341; 
on  the  incubation  of  the  Lop- 
hobranchii,  342;  on  rivalry  in 
song-birds,  364;  on  the  trachea  of 
the  swan,  370;  on  the  moulting 
of  the  Anatidse,  393;  on  the 
young  of  the  waders,  481. 

Yellow  fever,  immunity  of  ne- 
groes and  mulattoes  from,  188. 

Youatt,  Mr.,  on  the  development 
of  the  horns  in  cattle,  230. 

Yura-caras,  their  notions  of 
beauty,  574. 


Z 

Zebra,  rejection  of  an  ass  by  a 
female,  535;  stripes  of  the,  540. 

Zebus,  humps  of,  528. 

Zigzags,  prevalence  of,  as  "orna- 
ments,  175, 

Zincke,  Mr.,  on  European  emigra- 
tion to  America,  138. 

Zootoca  vivipara,  sexual  differ- 
ence in  the  color  of,  353. 

Zouteveen,  Dr.,  polydactylism,  35; 
proportion  of  sexes  at  Cape  of 
Good  Hope,  237;  spiders  at- 
tracted by  music,  270;  on  sounds 
produced  by  fish,  343. 

Ts&Ase,  coloration  of  the,  311. 


J 


Cbaiujos  Dah^ul 


DATE  DUE 

FFB  1  3  2 

]07 

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BOSTON  COLLEGE 


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